John K. B. Ford

Dialect change in resident killer whales: implications for vocal learning and cultural transmission

Variation in vocal signals among populations and social groups of animals provides opportunities for the study of the mechanisms of behavioural change and their importance in generating and maintaining behavioural variation. We analysed two call types made by two matrilineal social groups of resident killer whales, Orcinus orca, over 12-13 years. We used a neural network-based index of acoustic similarity to identify mechanisms of call differentiation. A test for structural modification of the calls detected significant changes in one call type in both groups, but not in the other. For the modified call type, the rate of divergence between the two groups was significantly lower than the rate of modification within either group showing that calls were modified in a similar fashion in the two groups. An analysis of structural parameters detected no strong directionality in the change. The pattern of call modification could have been caused by maturational changes to the calls or, if killer whale dialects are learned behavioural traits, cultural drift in the structure of the calls together with horizontal transmission of modifications between the two groups. Such vocal matching between members of different matrilines would suggest that vocal learning is not limited to vertical transmission from mother to offspring, which has important implications for models of gene-culture coevolution. Copyright 2000 The Association for the Study of Animal Behaviour.

Cultural transmission within maternal lineages: vocal clans in resident killer whales in southern Alaska

Cultural lineages are based on learned social traditions that are stable for several generations. When cultural lineages also reflect common ancestry and/or are shared by individuals that live together they are called clans. The existence of clans among killer whales has been previously proposed but has not been confirmed. Here, we show that clans exist among resident type killer whales, Orcinus orca, in southern Alaska. Resident killer whales live in stable matrilines from which emigration of either sex has not been observed. Matrilines that associate regularly (50% observation time) are called pods. Pods are believed to consist of closely related matrilines and share a unique repertoire of discrete call types. Pods that share parts of their repertoire form what Ford (1991, Canadian Journal of Zoology, 69, 1454–1483) called an acoustic clan. Here, we identified discrete call types of seven pods from southern Alaska, using a method based on human discrimination of distinct aural and visual (spectrogram) differences. Mitochondrial DNA of members of each pod was also analysed. The repertoires of the seven pods were compared and two acoustically distinct groups of pods were identified. Each group was monomorphic for a different mitochondrial D-loop haplotype. Nevertheless, pods from different clans associated frequently. It thus appears that the acoustic similarities within groups, which we presume to be cultural, reflect common ancestry, and that these groups therefore meet the above definition of clans. We also argue that a combination of cultural drift and selection are the main mechanisms for the maintenance of clans.

Selective habituation shapes acoustic predator recognition in harbour seals

Predation is a major force in shaping the behaviour of animals, so that precise identification of predators will confer substantial selective advantages on animals that serve as food to others. Because experience with a predator can be lethal, early researchers studying birds suggested that predator recognition does not require learning. However, a predator image that can be modified by learning and experience will be advantageous in situations where cues associated with the predator are highly variable or change over time. In this study, we investigated the response of harbour seals (Phoca vitulina) to the underwater calls of different populations of killer whales (Orcinus orca). We found that the seals responded strongly to the calls of mammal-eating killer whales and unfamiliar fish-eating killer whales but not to the familiar calls of the local fish-eating population. This demonstrates that wild harbour seals are capable of complex acoustic discrimination and that they modify their predator image by selectively habituating to the calls of harmless killer whales. Fear in these animals is therefore focused on local threats by learning and experience.

The vocal behaviour of mammal-eating killer whales: communicating with costly calls

The cost of vocal behaviour is usually expressed in energetic terms; however, many animals may pay additional costs when predators or potential prey eavesdrop on their vocal communication. The northeastern Pacific is home to two distinct ecotypes of killer whales, Orcinus orca, called residents and transients. Resident killer whales feed on fish, a prey with poor hearing abilities, whereas transient killer whales hunt marine mammals, which have sensitive underwater hearing within the frequency range of killer whale vocal communication. In this study, we investigated how the superior hearing ability of mammalian prey has shaped the vocal behaviour of the transient killer whale ecotype. We recorded pulsed calls and the associated behavioural context of groups of transient and resident killer whales in British Columbia and southeastern Alaska. Transient killer whales produced pulsed calls significantly less frequently than residents. Transient killer whales only showed significant amounts of vocal behaviour after a marine mammal kill or when the whales were displaying surface-active behaviour. Vocal activity of transients increased after a successful attack on a marine mammal. Since marine mammals are able to detect killer whale pulsed calls and respond with antipredator behaviour, the reduced vocal activity of transients is probably due to a greater cost for calling in this ecotype resulting from eavesdropping by potential prey. The increase in vocal behaviour after a successful attack may represent food calling (informing other animals in the area about the presence of food), but is more likely to reflect an increase in social interactions during feeding and/or the fact that the cost for vocal behaviour is comparatively low after a successful attack.

Linking killer whale survival and prey abundance: food limitation in the oceans' apex predator?

Killer whales (Orcinus orca) are large predators that occupy the top trophic position in the worlds oceans and as such may have important roles in marine ecosystem dynamics. Although the possible top-down effects of killer whale predation on populations of their prey have received much recent attention, little is known of how the abundance of these predators may be limited by bottom-up processes. Here we show, using 25 years of demographic data from two populations of fish-eating killer whales in the northeastern Pacific Ocean, that population trends are driven largely by changes in survival, and that survival rates are strongly correlated with the availability of their principal prey species, Chinook salmon (Oncorhynchus tshawytscha). Our results suggest that, although these killer whales may consume a variety of fish species, they are highly specialized and dependent on this single salmonid species to an extent that it is a limiting factor in their population dynamics. Other ecologically specialized killer whale populations may be similarly constrained to a narrow range of prey species by culturally inherited foraging strategies, and thus are limited in their ability to adapt rapidly to changing prey availability.

Adaptive Prolonged Postreproductive Life Span in Killer Whales

Killer whale mothers continue to help their adult male offspring to survive long after ceasing reproduction. Prolonged life after reproduction is difficult to explain evolutionarily unless it arises as a physiological side effect of increased longevity or it benefits related individuals (i.e., increases inclusive fitness). There is little evidence that postreproductive life spans are adaptive in nonhuman animals. By using multigenerational records for two killer whale (Orcinus orca) populations in which females can live for decades after their final parturition, we show that postreproductive mothers increase the survival of offspring, particularly their older male offspring. This finding may explain why female killer whales have evolved the longest postreproductive life span of all nonhuman animals.

Stability and group specificity of stereotyped whistles in resident killer whales, Orcinus orca, off British Columbia

Resident killer whales off British Columbia form four acoustically distinct clans, each with a unique dialect of discrete pulsed calls. Three clans belong to the northern and one to the southern community. Resident killer whales also produce tonal whistles, which play an important role in close-range communication within the northern community. However, there has been no comparative analysis of repertoires of whistles across clans. We investigated the structural characteristics, stability and group specificity of whistles in resident killer whales off British Columbia. Acoustic recordings and behavioural observations were made between 1978 and 2003. Whistles were classified spectrographically and additional observers were used to confirm our classification. Whistles were compared across clans using discriminant function analysis. We found 11 types of stereotyped whistles in the northern and four in the southern community with some of the whistle types being stable over at least 13 years. In northern residents, 10 of the 11 whistle types were structurally identical in two of the three acoustic clans, whereas the whistle types of southern residents differed clearly from those of the northern residents. Our study shows that killer whales that have no overlap in their call repertoire use essentially the same set of stereotyped whistles. Shared stereotyped whistles might provide a community-level means of recognition that facilitates association and affiliation of members of different clans, which otherwise use distinct signals. We further suggest that vocal learning between groups plays an important role in the transmission of whistle types.

Killer Whale: Orcinus orca

Summary The killer whale, the ocean’s apex predator, has a cosmopolitan range through all the worlds oceans and most seas. It is the largest member of the family Delphinidae and has very distinctive black-and-white coloration. Only a single species, Orcinus orca , is currently recognized but it is probable that some of the genetically distinct forms found in different regions represent distinct species. Although a generalist as a species, different populations of killer whales usually have specialized foraging behavior and diets. These ecotypes often employ innovative cooperative hunting tactics to catch prey. Killer whales generally live in stable matrilineally structured groups from which dispersal of individuals may be rare on nonexistent. Killer whales produce highly structured, stereotyped calls that vary among populations and, often, among different maternal lineages. Once widely considered to be dangerous to humans or nuisances that interfered with fisheries, killer whales were widely persecuted. Today they are generally admired and protected, and are a focus of whale watching tourism in many regions. Availability of prey, acoustic disturbance, and contaminants are considered the primary anthropogenic threats to the species.

Characteristics of whistles from the acoustic repertoire of resident killer whales (Orcinus orca) off Vancouver Island, British Columbia

The acoustic repertoire of killer whales (Orcinus orca) consists of pulsed calls and tonal sounds, called whistles. Although previous studies gave information on whistle parameters, no study has presented a detailed quantitative characterization of whistles from wild killer whales. Thus an interpretation of possible functions of whistles in killer whale underwater communication has been impossible so far. In this study acoustic parameters of whistles from groups of individually known killer whales were measured. Observations in the field indicate that whistles are close-range signals. The majority of whistles (90%) were tones with several harmonics with the main energy concentrated in the fundamental. The remainder were tones with enhanced second or higher harmonics and tones without harmonics. Whistles had an average bandwidth of 4.5 kHz, an average dominant frequency of 8.3 kHz, and an average duration of 1.8 s. The number of frequency modulations per whistle ranged between 0 and 71. The study indicates that whistles in wild killer whales serve a different function than whistles of other delphinids. Their structure makes whistles of killer whales suitable to function as close-range motivational sounds.

On the communicative significance of whistles in wild killer whales (Orcinus orca)

Abstract. Killer whales (Orcinus orca) use pulsed calls and whistles in underwater communication. Unlike pulsed calls, whistles have received little study and thus their function is poorly known. In this study, whistle activities of groups of individually known killer whales were compared quantitatively across behavioural categories. Acoustic recordings and simultaneous behavioural observations were made of northern resident killer whales off Vancouver Island in 1996 and 1997. Whistles were produced at greater rates than discrete calls during close-range behavioural activities than during long-range activities. They were the predominant sound-type recorded during socializing. The number of whistles per animal per minute was significantly higher during close-range behavioural activities than during long-range activities. Evidently, whistles play an important role in the close-range acoustic communication in northern resident killer whales.