Luke Rendell

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.

Why copy others? Insights from the social learning strategies tournament.

It Pays to Be a Copy Cat Does it pay to copy what others do? Rendell et al. (p. 208) elected to copy Robert Axelrods 1979 tournament in which strategies for playing the iterated prisoners dilemma game were pitted against each other until an overall winner emerged—the tit-for-tat strategy. In the 2008 tournament, 100 social learning strategies designed to cope with a changing environment competed against each other; the winning strategy involved sampling the behaviors of other players periodically, rather than exploring the environment alone. Learning from what others do is more efficient than learning all on one’s own. Social learning (learning through observation or interaction with other individuals) is widespread in nature and is central to the remarkable success of humanity, yet it remains unclear why copying is profitable and how to copy most effectively. To address these questions, we organized a computer tournament in which entrants submitted strategies specifying how to use social learning and its asocial alternative (for example, trial-and-error learning) to acquire adaptive behavior in a complex environment. Most current theory predicts the emergence of mixed strategies that rely on some combination of the two types of learning. In the tournament, however, strategies that relied heavily on social learning were found to be remarkably successful, even when asocial information was no more costly than social information. Social learning proved advantageous because individuals frequently demonstrated the highest-payoff behavior in their repertoire, inadvertently filtering information for copiers. The winning strategy (discountmachine) relied nearly exclusively on social learning and weighted information according to the time since acquisition.

Cognitive culture: theoretical and empirical insights into social learning strategies

Research into social learning (learning from others) has expanded significantly in recent years, not least because of productive interactions between theoretical and empirical approaches. This has been coupled with a new emphasis on learning strategies, which places social learning within a cognitive decision-making framework. Understanding when, how and why individuals learn from others is a significant challenge, but one that is critical to numerous fields in multiple academic disciplines, including the study of social cognition.

Network-Based Diffusion Analysis Reveals Cultural Transmission of Lobtail Feeding in Humpback Whales

Animal Culture Cultural transmission of information occurs when individuals learn from others with more experience or when individuals come to accept particular modes of behavior as the local norm. Such information transfer can be expected in highly social or long-lived species where contact and time for learning are maximized and are seen in humans (see the Perspective by de Waal). Using a network-based diffusion analysis on a long-term data set that includes tens of thousands of observations of individual humpback whales, Allen et al. (p. 485) show that an innovative feeding behavior has spread through social transmission since it first emerged in a single individual in 1980. The “lobtail” feeding has passed among associating individuals for more than three decades. Van de Waal et al. (p. 483), on the other hand, used a controlled experimental approach in vervet monkeys to show that individuals learn what to eat from more experienced individuals within their social group. Not only did young animals learn from observing older animals, but immigrating males switched their food preference to that of their new group. Whales have learned from their peers how to use their tails to herd prey. [Also see Perspective by de Waal] We used network-based diffusion analysis to reveal the cultural spread of a naturally occurring foraging innovation, lobtail feeding, through a population of humpback whales (Megaptera novaeangliae) over a period of 27 years. Support for models with a social transmission component was 6 to 23 orders of magnitude greater than for models without. The spatial and temporal distribution of sand lance, a prey species, was also important in predicting the rate of acquisition. Our results, coupled with existing knowledge about song traditions, show that this species can maintain multiple independently evolving traditions in its populations. These insights strengthen the case that cetaceans represent a peak in the evolution of nonhuman culture, independent of the primate lineage.

Vocal clans in sperm whales (Physeter macrocephalus).

Cultural transmission may be a significant source of variation in the behaviour of whales and dolphins, especially as regards their vocal signals. We studied variation in the vocal output of ‘codas’ by sperm whale social groups. Codas are patterns of clicks used by female sperm whales in social circumstances. The coda repertoires of all known social units (n = 18, each consisting of about 11 females and immatures with long–term relationships) and 61 out of 64 groups (about two social units moving together for periods of days) that were recorded in the South Pacific and Caribbean between 1985 and 2000 can be reliably allocated into six acoustic ‘clans’, five in the Pacific and one in the Caribbean. Clans have ranges that span thousands of kilometres, are sympatric, contain many thousands of whales and most probably result from cultural transmission of vocal patterns. Units seem to form groups preferentially with other units of their own clan. We suggest that this is a rare example of sympatric cultural variation on an oceanic scale. Culture may thus be a more important determinant of sperm whale population structure than genes or geography, a finding that has major implications for our understanding of the species’ behavioural and population biology.

Experimental evidence for the co-evolution of hominin tool-making teaching and language

Hominin reliance on Oldowan stone tools – which appear from 2.5mya and are believed to have been socially transmitted – has been hypothesised to have led to the evolution of teaching and language. Here we present an experiment investigating the efficacy of transmission of Oldowan tool-making skills along chains of adult human participants (N=184) using 5 different transmission mechanisms. Across six measures, transmission improves with teaching, and particularly with language, but not with imitation or emulation. Our results support the hypothesis that hominin reliance on stone tool-making generated selection for teaching and language and imply that (i) low-fidelity social transmission, such as imitation/emulation, may have contributed to the ~700,000 year stasis of the Oldowan technocomplex, and (ii) teaching or proto-language may have been pre-requisites for the appearance of Acheulean technology. This work supports a gradual evolution of language, with simple symbolic communication preceding behavioural modernity by hundreds of thousands of years.

Lessons from animal teaching

Many species are known to acquire valuable life skills and information from others, but until recently it was widely believed that animals did not actively facilitate learning in others. Teaching was regarded as a uniquely human faculty. However, recent studies suggest that teaching might be more common in animals than previously thought. Teaching is present in bees, ants, babblers, meerkats and other carnivores but is absent in chimpanzees, a bizarre taxonomic distribution that makes sense if teaching is treated as a form of altruism. Drawing on both mechanistic and functional arguments, we integrate teaching with the broader field of animal social learning, and show how this aids understanding of how and why teaching evolved, and the diversity of teaching mechanisms.

Rogers' paradox recast and resolved: population structure and the evolution of social learning strategies.

We explore the evolution of reliance on social and asocial learning using a spatially explicit stochastic model. Our analysis considers the relative merits of four evolved strategies, two pure strategies (asocial and social learning) and two conditional strategies (the “critical social learner,” which learns asocially only when copying fails, and the “conditional social learner,” which copies only when asocial learning fails). We find that spatial structure generates outcomes that do not always conform to the finding of earlier theoretical analyses that social learning does not enhance average individual fitness at equilibrium (Rogers’ paradox). Although we describe circumstances under which the strategy of pure social learning increases the average fitness of individuals, we find that spatial structure introduces a new paradox, which is that social learning can spread even when it decreases the average fitness of individuals below that of asocial learners. We also show that the critical social learner and conditional social learner both provide solutions to the aforementioned paradoxes, although we find some conditions in which pure (random) social learning out‐competes both conditional strategies. Finally, we consider the relative merits of critical and conditional social learning under various conditions.

Female philopatry in coastal basins and male dispersion across the North Atlantic in a highly mobile marine species, the sperm whale (Physeter macrocephalus).

The mechanisms that determine population structure in highly mobile marine species are poorly understood, but useful towards understanding the evolution of diversity, and essential for effective conservation and management. In this study, we compare putative sperm whale populations located in the Gulf of Mexico, western North Atlantic, Mediterranean Sea and North Sea using mtDNA control region sequence data and 16 polymorphic microsatellite loci. The Gulf of Mexico, western North Atlantic and North Sea populations each possessed similar low levels of haplotype and nucleotide diversity at the mtDNA locus, while the Mediterranean Sea population showed no detectable mtDNA diversity. Mitochondrial DNA results showed significant differentiation between all populations, while microsatellites showed significant differentiation only for comparisons with the Mediterranean Sea, and at a much lower level than seen for mtDNA. Samples from either side of the North Atlantic in coastal waters showed no differentiation for mtDNA, while North Atlantic samples from just outside the Gulf of Mexico (the western North Atlantic sample) were highly differentiated from samples within the Gulf at this locus. Our analyses indicate a previously unknown fidelity of females to coastal basins either side of the North Atlantic, and suggest the movement of males among these populations for breeding.

A REVIEW OF FREQUENCY AND TIME PARAMETERS OF CETACEAN TONAL CALLS

ABSTRACT Properties of the tonal calls of cetaceans are summarised and compared at the species level. Statistics are presented relating to start, end, minimum, maximum and centre frequencies, duration and number of inflections, together with information about recordings (location, number encounters/groups, length of recordings). Evidence of a linear relation between body-size and wavelength is given for odontocetes (R2 between 0.68 and 0.93, depending on frequency variable and least-squares method) but the relationship does not appear to fit well for mysticetes (R2 = 0.64). Results of preliminary investigations into acoustic discrimination of species by multivariate methods are also presented: a simple classifier for 10 species based on results derived from the literature gave a correct classification rate of 28% when tested with independent data. Some methodological recommendations for future descriptive work on cetacean acoustics are made.