Janet Mann

Cetacean societies : field studies of dolphins and whales

Long-lived, slow to reproduce, and often hidden beneath the waters surface, whales and dolphins (cetaceans) have remained elusive subjects for scientific study even though they have fascinated humans for centuries. Until recently, much of what we knew about cetaceans came from commercial sources such as whalers and trainers for dolphin acts. Innovative research methods and persistent efforts, however, have begun to penetrate the depths to reveal tantalizing glimpses of the lives of these mammals in their natural habitats. This book presents a comprehensive synthesis and review of these studies. Groups of chapters focus on the history of cetacean behavioural research and methodology; state-of-the-art reviews of information on four of the most-studied species: bottlenose dolphins, killer whales, sperm whales and humpback whales; and summaries of major topics, including group living, male and female reproductive strategies, communication, and conservation drawn from comparative research on a wide range of species. Written by cetacean scientists, this volume should be of benefit to students of cetology and researchers in other areas of behavioral and conservation ecology, as well as anyone with a serious interest in the world of whales and dolphins.

Social evolution in toothed whales

Two contrasting results emerge from comparisons of the social systems of several odontocetes with terrestrial mammals. Researchers have identified remarkable convergence in prominent features of the social systems of odontocetes such as the sperm whale and bottlenose dolphin with a few well-known terrestrial mammals such as the elephant and chimpanzee. In contrast, studies on killer whales and Bairds beaked whale reveal novel social solutions to aquatic living. The combination of convergent and novel features in odontocete social systems promise a more general understanding of the ecological determinants of social systems in both terrestrial and aquatic habitats, as well as the relationship between relative brain size and social evolution.

Use of signature whistles during separations and reunions by wild bottlenose dolphin mothers and infants

We examine the contexts and patterns of “signature” whistle production by wild bottlenose dolphin mother-infant pairs (Tursiops spp.) to gain insight into the functional significance of whistles. Results are based on focal observations and simultaneous recordings of underwater vocalizations. Whistles occur primarily when mother-infant pairs are separated, and the probability of whistles increases with distance of separation. The timing of whistles during separations varies, but whistles tend to be produced in repetitive series and are generally concentrated toward the later stages of the separation, i.e., during the process of reunion. Although we focused on infants, mothers do not appear to whistle during separations as frequently as infants. Infant whistles may function to facilitate reunions by conveying information to the mother concerning the infants motivation to reunite and/or its location. Infant whistles could induce a cooperative response from the mother including approach, slowing to allow the infant to catch up or whistling. Highly individualized signature whistles may be particularly useful in a fission-fusion society in which individuals (mothers and infants as well as adults) join and leave temporary parties in a fluid manner, yet maintain consistent, long-term associations with particular individuals.

Aggression in bottlenose dolphins: Evidence for sexual coercion, male-male competition, and female tolerance through analysis of tooth-rake marks and behaviour

Aggressive behaviour is rarely observed, but may have a large impact on the social structure, relationships and interactions in animal societies. Long-term behavioural study of Indian Ocean bottlenose dolphins in Shark Bay, Australia, suggests that males are more aggressive than females, and use sexual coercion during the breeding season, but age and sex-specific patterns of aggression have not been well documented. We analyzed tooth rake marks, an indirect measure of received conspecific aggression, to determine such patterns by age, sex, and adult female reproductive state. Photographs of 224 Shark Bay bottlenose dolphins were examined for tooth rakes and each rake was categorized as new (broken skin), obvious (white rake lines that are clearly visible) or faint (faint evidence of rakes). Rake lines were also coded by each body section visible in the photograph. Cycling females (those that became pregnant within 6 mos. of the photograph date) were significantly more likely to have new tooth rakes than non-cycling females (pregnant or with a dependent calf 2 years apart) suggests that the marks are not cumulative. Overall, 83% of the population has tooth rake marks, suggesting that agonistic interactions occur for most individuals at least every two years. Analysis of gonistic interactions for 55 adult focal females (observed for 1960 h) and their 88 calves (observed for 1876 h) show that female aggression is extremely rare (occurring every 490 h), but females receive aggression more often (every 61 h), and 84% of received aggression was from juvenile and adult males. Focal adult females were never observed acting aggressively towards juveniles or adults of either sex, suggesting that female bottlenose dolphins are highly tolerant. Of the four observed instances of adult female aggression, all were directed at their dependent offspring. Male calves are significantly more aggressive than female calves, and calves had higher rates of agonistic interactions than their mothers (every 18 h), even though calves had fewer tooth rakes than all other age classes. The patterns of tooth rake presence and prevalence likely result from sexual coercion of adult females by adult males and intra-sexual male competition.

Social and genetic interactions drive fitness variation in a free-living dolphin population

The evolutionary forces that drive fitness variation in species are of considerable interest. Despite this, the relative importance and interactions of genetic and social factors involved in the evolution of fitness traits in wild mammalian populations are largely unknown. To date, a few studies have demonstrated that fitness might be influenced by either social factors or genes in natural populations, but none have explored how the combined effect of social and genetic parameters might interact to influence fitness. Drawing from a long-term study of wild bottlenose dolphins in the eastern gulf of Shark Bay, Western Australia, we present a unique approach to understanding these interactions. Our study shows that female calving success depends on both genetic inheritance and social bonds. Moreover, we demonstrate that interactions between social and genetic factors also influence female fitness. Therefore, our study represents a major methodological advance, and provides critical insights into the interplay of genetic and social parameters of fitness.

PATTERNS OF FEMALE ATTRACTIVENESS IN INDIAN OCEAN BOTTLENOSE DOLPHINS

Hormonal profiles of captive individuals show that bottlenose dolphins (Tursiops spp.) are seasonally polyoestrous, but little is known of reproductive behaviour among free-ranging bottlenose dolphins. In Shark Bay, Western Australia, we have documented for the first time patterns of female attractiveness that may correspond to multiple oestrous cycles. Male bottlenose dolphins in stable alliances of 2-3 individuals form temporary consortships with individual females. Consortships often are established and maintained by aggressive herding. Consortships are associated with reproduction and are a useful measure of a females attractiveness. Following reproduction, females may become attractive to males when their surviving calf is about 2-2.5 years old or within 1-2 weeks of losing an infant. Individual females are attractive to males for variable periods extending over a number of months, both within and outside of the main breeding season. The duration of attractive periods is greater during breeding season months than during the preceding months. Males sometimes are attracted to females for periods exceeding the reported duration of rising estrogen levels during the follicular stage of the oestrous cycle. Males occasionally have consorted or otherwise been attracted to females in several unusual contexts, including late pregnancy, the first two weeks after parturition, and the day after the loss of a nursing infant. Individual females were consorted by up to 13 males during the season they conceived, supporting predictions of a promiscuous mating system in bottlenose dolphins. Thus, consorting is a strategy by males to monopolize females, but not a completely successful one. Multiple cycling by female bottlenose dolphins may be a strategy to avoid being monopolized by particular males. Given the duration and agonistic nature of many consortships, the benefits to females of such a costly strategy are not obvious. Multiple cycling may reduce the risk of infanticide by males or allow females to mate with preferred males after being monopolized by less desirable males.

Contrasting relatedness patterns in bottlenose dolphins (Tursiops sp.) with different alliance strategies

Male bottlenose dolphins (Tursiops sp.) in Shark Bay have one of the most complex male societies outside humans. Two broad mating strategies have been identified in males. In the first strategy, there are two types of alliances: stable‘first–order’ pairs and trios that herd individual females in reproductive condition, and ‘second–order’ teams of two first–order alliances (five or six individuals) that join forces against rivals in contests for females. In the alternative strategy, a ‘super–alliance’ of ca. 14 individuals, males form pairs or trios to herd females, but in contrast to the stable alliances, these pairs and trios are highly labile. Here, we show that males in stable first–order alliances and the derived second–order alliances are often strongly related, so that they may gain inclusive fitness benefits from alliance membership. By contrast, members of the super–alliance are no more closely related than expected by chance. Further, the strength of the association of alliance partners within the super–alliance, as measured by an index of joint participation in consorting a female, was not correlated with their genetic relatedness. Thus, within one population and one sex, it appears that there may be simultaneous operation of more than one mode of group formation.

'O father: where art thou?'--Paternity assessment in an open fission-fusion society of wild bottlenose dolphins (Tursiops sp.) in Shark Bay, Western Australia.

Sexually mature male bottlenose dolphins in Shark Bay cooperate by pursuing distinct alliance strategies to monopolize females in reproductive condition. We present the results of a comprehensive study in a wild cetacean population to test whether male alliance membership is a prerequisite for reproductive success. We compared two methods for inferring paternity: both calculate a likelihood ratio, called the paternity index, between two opposing hypotheses, but they differ in the way that significance is applied to the data. The first method, a Bayesian approach commonly used in human paternity testing, appeared to be overly conservative for our data set, but would be less susceptible to assumptions if a larger number of microsatellite loci had been used. Using the second approach, the computer program cervus 2.0, we successfully assigned 11 paternities to nine males, and 17 paternities to 14 out of 139 sexually mature males at 95% and 80% confidence levels, respectively. It appears that being a member of a bottlenose dolphin alliance is not a prerequisite for paternity: two paternities were obtained by juvenile males (one at the 95%, the other at the 80% confidence level), suggesting that young males without alliance partners pursue different mating tactics to adults. Likelihood analyses showed that these two juvenile males were significantly more likely to be the true father of the offspring than to be their half‐sibling (P < 0.05). Using paternity data at an 80% confidence level, we could show that reproductive success was significantly skewed within at least some stable first‐order alliances (P < 0.01). Interestingly, there is powerful evidence that one mating was incestuous, with one calf apparently fathered by its mothers father (P < 0.01). Our study suggests that the reproductive success of both allied males, and of nonallied juveniles, needs to be incorporated into an adaptive framework that seeks to explain alliance formation in male bottlenose dolphins.

Why Do Dolphins Carry Sponges

Tool use is rare in wild animals, but of widespread interest because of its relationship to animal cognition, social learning and culture. Despite such attention, quantifying the costs and benefits of tool use has been difficult, largely because if tool use occurs, all population members typically exhibit the behavior. In Shark Bay, Australia, only a subset of the bottlenose dolphin population uses marine sponges as tools, providing an opportunity to assess both proximate and ultimate costs and benefits and document patterns of transmission. We compared sponge-carrying (sponger) females to non-sponge-carrying (non-sponger) females and show that spongers were more solitary, spent more time in deep water channel habitats, dived for longer durations, and devoted more time to foraging than non-spongers; and, even with these potential proximate costs, calving success of sponger females was not significantly different from non-spongers. We also show a clear female-bias in the ontogeny of sponging. With a solitary lifestyle, specialization, and high foraging demands, spongers used tools more than any non-human animal. We suggest that the ecological, social, and developmental mechanisms involved likely (1) help explain the high intrapopulation variation in female behaviour, (2) indicate tradeoffs (e.g., time allocation) between ecological and social factors and, (3) constrain the spread of this innovation to primarily vertical transmission.

Developmental evidence for foraging traditions in wild bottlenose dolphins

Patterns of social learning in bottlenose dolphins (Tursiops spp.) may help explain behavioural variants and selection pressures favouring cultural evolution, but evidence for social transmission derived from field observations is controversial. The dolphins of Shark Bay, Australia are known for diverse, individually specific foraging behaviours, including tool use with marine sponges. We examined the relative contributions of habitat, maternal foraging and foraging behaviours of associates to the use of seven foraging tactics by bottlenose dolphin calves (Tursiops sp.). Three tactics were predicted by maternal foraging, one was predicted by water depth, one was weakly predicted by several factors, and two tactics were not predicted by any variable. Our findings provide support for the social learning of foraging behaviours in wild dolphins, illustrate the diverse pathways of foraging development, and offer insight into conditions that are likely to favour reliance on social information The evidence for vertical social learning (mother-to-offspring) indicates the dominance of a ‘do what mother does’ strategy, rather than copying the foraging behaviour of associates. However, since not all foraging behaviours were predicted by whether a calfs mother used them, dolphins may only use social information in some contexts, perhaps for more difficult tactics. This study provides unprecedented support for socially learned foraging tactics in wild dolphins by simultaneously addressing multiple factors during behavioural development, and thus illustrates the benefits of using multivariable techniques on ontogenetic data to identify social learning in wild animals.