Characterizing the socially transmitted foraging tactic “sponging” by bottlenose dolphins (Tursiops sp.) in the western gulf of Shark Bay,Western Australia

Individual foraging tactics are widespread in animals and have ecological and evolutionary implications. Indo‐Pacific bottlenose dolphins (Tursiops sp.) in Shark Bay, Western Australia, exhibit a foraging tactic involving tool use, called “sponging.” Sponging is vertically, socially transmitted through the matriline and, to date, has been described in detail in the eastern gulf of Shark Bay (ESB). Here, we characterize sponging in the western gulf of Shark Bay (WSB), in which a different matriline engages in the behavior. We identified 40 individual “spongers” in 9 mo of boat‐based surveys over three field seasons. As is the case in ESB, the majority of WSB spongers was female and engaged in sponging in deep channel habitats. In contrast to ESB, however, there was no difference in the number of associates between spongers and nonspongers in WSB, and activity budgets differed between spongers and deep‐water nonspongers; spongers foraged more frequently and rested less than nonspongers. Group sizes in deep channel habitat, where sponging was prevalent, were typically larger than those in shallow habitat, except for foraging, perhaps indicative of higher predator abundance and/or scattered prey distribution in deep‐water habitat. This research improves our understanding of within‐population foraging variations in 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.     

Cultural transmission of tool use combined with habitat specializations leads to fine-scale genetic structure in bottlenose dolphins

Socially learned behaviours leading to genetic population structure have rarely been described outside humans. Here, we provide evidence of fine-scale genetic structure that has probably arisen based on socially transmitted behaviours in bottlenose dolphins (Tursiops sp.) in western Shark Bay, Western Australia. We argue that vertical social transmission in different habitats has led to significant geographical genetic structure of mitochondrial DNA (mtDNA) haplotypes. Dolphins with mtDNA haplotypes E or F are found predominantly in deep (more than 10 m) channel habitat, while dolphins with a third haplotype (H) are found predominantly in shallow habitat (less than 10 m), indicating a strong haplotype–habitat correlation. Some dolphins in the deep habitat engage in a foraging strategy using tools. These ‘sponging’ dolphins are members of one matriline, carrying haplotype E. This pattern is consistent with what had been demonstrated previously at another research site in Shark Bay, where vertical social transmission of sponging had been shown using multiple lines of evidence. Using an individual-based model, we found support that in western Shark Bay, socially transmitted specializations may have led to the observed genetic structure. The reported genetic structure appears to present an example of cultural hitchhiking of mtDNA haplotypes on socially transmitted foraging strategies, suggesting that, as in humans, genetic structure can be shaped through cultural transmission.     

Sponge Carrying by Dolphins (Delphinidae,Tursiops sp.): A Foraging Specialization Involving Tool Use?

During long-term research on bottlenose dolphins (Tursiops sp.) in Shark Bay, Western Australia, several individuals were observed carrying sponges, Echinodictyum mesenterinum, on their rostra. Over multiple years, five regularly sighted individuals were usually carrying sponges when encountered (67–100% of encounters). Four additional regularly sighted individuals were observed with sponges just one time each. All five individuals that routinely carried sponges were female. Two of the anomalous, one-time carriers were female, one was likely female, and one was male. Most observations of sponge carrying occurred within a restricted area, a relatively deep water channel (8–10 m deep). Surface observations of sponge carrying, including focal animal observations, revealed a stereotyped surfacing and diving pattern, and occasional indications of prey consumption. Three hypotheses are considered regarding the function of sponge carrying: 1. dolphins were playing with the sponges; 2. the sponges contain some compound of use to the dolphins (e.g. for medicinal purposes); and 3. the sponges were used as a tool to aid in foraging. The foraging tool hypothesis is best supported, but the exact manner in which sponges are used remains to be discovered. Sponge carrying is a behavioural specialization, probably involving foraging, and regularly engaged in by only a small proportion of female dolphins in Shark Bay.     

Where to catch a fish? The influence of foraging tactics on the ecology of bottlenose dolphins (Tursiops truncatus) in Florida Bay,Florida

Observations of bottlenose dolphins ( Tursiops truncatus ) in Florida Bay, Florida, between 2002 and 2005 revealed the use of three distinct foraging tactics. The goal of this study was to identify ecological correlates with tactic use and describe the impact of foraging specializations on the overall habitat use and distribution patterns of this dolphin population. Foraging tactics showed strong association with contrasting environmental characteristics, primarily depth. Locations of two of these tactic groups were spatially repulsed. Analyses of sighting histories of individual dolphins observed at foraging events determined that dolphins which employed one tactic never employed the other, and vice versa . Although bottlenose dolphins have plastic foraging behaviors, dolphins in Florida Bay appear to specialize in one tactic and subsequently limit their overall distribution patterns to coincide with habitats that facilitate success using that foraging tactic. This study demonstrates how foraging behavior can be an ecological determinant of overall dolphin habitat use patterns and works to create spatial structure within a population due to consistent mapping of tactics onto environmental variation. These foraging specializations potentially impact the social and demographic patterns of this dolphin population. The possible evolutionary mechanisms behind this intraspecific variation, including resource limitation and social learning, are considered.     

The ecological conditions that favor tool use and innovation in wild bottlenose dolphins (Tursiops sp.)

Dolphins are well known for their exquisite echolocation abilities, which enable them to detect and discriminate prey species and even locate buried prey. While these skills are widely used during foraging, some dolphins use tools to locate and extract prey. In the only known case of tool use in free-ranging cetaceans, a subset of bottlenose dolphins (Tursiops sp.) in Shark Bay, Western Australia habitually employs marine basket sponge tools to locate and ferret prey from the seafloor. While it is clear that sponges protect dolphins' rostra while searching for prey, it is still not known why dolphins probe the substrate at all instead of merely echolocating for buried prey as documented at other sites. By 'sponge foraging' ourselves, we show that these dolphins target prey that both lack swimbladders and burrow in a rubble-littered substrate. Delphinid echolocation and vision are critical for hunting but less effective on such prey. Consequently, if dolphins are to access this burrowing, swimbladderless prey, they must probe the seafloor and in turn benefit from using protective sponges. We suggest that these tools have allowed sponge foraging dolphins to exploit an empty niche inaccessible to their non-tool-using counterparts. Our study identifies the underlying ecological basis of dolphin tool use and strengthens our understanding of the conditions that favor tool use and innovation in the wild.     

Evidence for Distinct Coastal and Offshore Communities of Bottlenose Dolphins in the North East Atlantic

Bottlenose dolphin stock structure in the northeast Atlantic remains poorly understood. However, fine scale photo-id data have shown that populations can comprise multiple overlapping social communities. These social communities form structural elements of bottlenose dolphin (Tursiops truncatus) populations, reflecting specific ecological and behavioural adaptations to local habitats. We investigated the social structure of bottlenose dolphins in the waters of northwest Ireland and present evidence for distinct inshore and offshore social communities. Individuals of the inshore community had a coastal distribution restricted to waters within 3 km from shore. These animals exhibited a cohesive, fission-fusion social organisation, with repeated resightings within the research area, within a larger coastal home range. The offshore community comprised one or more distinct groups, found significantly further offshore (>4 km) than the inshore animals. In addition, dorsal fin scarring patterns differed significantly between inshore and offshore communities with individuals of the offshore community having more distinctly marked dorsal fins. Specifically, almost half of the individuals in the offshore community (48%) had characteristic stereotyped damage to the tip of the dorsal fin, rarely recorded in the inshore community (7%). We propose that this characteristic is likely due to interactions with pelagic fisheries. Social segregation and scarring differences found here indicate that the distinct communities are likely to be spatially and behaviourally segregated. Together with recent genetic evidence of distinct offshore and coastal population structures, this provides evidence for bottlenose dolphin inshore/offshore community differentiation in the northeast Atlantic. We recommend that social communities should be considered as fundamental units for the management and conservation of bottlenose dolphins and their habitat specialisations.     

Shark bite injuries on three inshore dolphin species in tropical northwestern Australia

Predation risk has a profound influence on the behavior of marine mammals, affecting grouping patterns and habitat use. Dolphins frequently bear evidence of shark bites, which can provide an indirect measure of predation pressure. Using photo‐identification data, we investigated the prevalence of shark bites on three sympatric species of inshore dolphin, the Australian snubfin (Orcaella heinsohni), Australian humpback (Sousa sahulensis), and Indo‐Pacific bottlenose dolphin (Tursiops aduncus), among four study sites in northwestern Australia. Bite prevalence varied markedly between species, with 72% of snubfin, 46% of humpback, and 18% of bottlenose dolphins exhibiting evidence of shark bites. Binomial logistic regression confirmed a high likelihood of bite presence on snubfin dolphins, and at one particular site for snubfin and bottlenose dolphins. The prevalence of tiger shark (Galeocerdo cuvier) bites on snubfin dolphins was high, and bites attributed to other carcharhinid sharks were observed on all species. While acknowledging methodological differences with other studies, the prevalence of shark bites on snubfin dolphins is among the highest reported for any dolphins, suggesting predation risk represents an important but varying influence thereon. This study provides a baseline for future investigations into the affect of predation risk on the behavioral ecology of these sympatric species.     

Food-related bray calls in wild bottlenose dolphins (Tursiops truncatus)

Because cetaceans are difficult to study in the wild, little is known about how they use their sounds in their natural environment. Only the recent development of passive acoustic localization systems has enabled observations of the communication behaviour of individuals for correlation with their surface behaviour. Using such a system, I show that bottlenose dolphins in the Moray Firth, Scotland, produce low-frequency bray calls which are clearly correlated with feeding on salmonids. The production of these calls is followed by fast approaches by conspecifics in the area. In animals which use sound as a foraging tool, it is difficult to distinguish between food calls which have evolved because of their role in attracting conspecifics, and food manipulation or searching calls which may attract conspecifics as a by-product. However, the low-frequency structure of the bottlenose dolphin bray suggests that it evolved because of a role in manipulating prey rather than in attracting conspecifics. This conclusion suggests that dolphins exploit the perceptual systems of their prey to facilitate capture.     

Spatial and Social Sexual Segregation Patterns in Indo-Pacific Bottlenose Dolphins (Tursiops aduncus)

Sexual segregation seems to be common in bottlenose dolphins, whereby males and females live in different pods that mix mainly for mating. Male dolphins often use aggressive behaviour to mate with females, while females with calves may have different activity and dietary requirements to males and different susceptibility to predation. We investigated the degree of spatial and social sexual segregation in Indo-Pacific bottlenose dolphins (Tursiops aduncus) in a subtropical estuary in Australia. Based on surveys completed over three years, dolphin groups were mostly mixed-sex or female. Mixed-sex groups were found in larger groups in mostly deeper water, whereas, female groups were foraging across all water depths in smaller groups. Aggressive coercive behaviour by males towards females was high, occurring mainly in deeper water, at higher tides, and outside the breeding season. Habitat use by female dolphin groups suggests that shallow tributaries may provide a sanctuary from aggressive males, access to suitable prey items and density for mothers and their calves, or a combination of these factors.     

A comparison of pectoral fin contact between two different wild dolphin populations

Contact behaviour involving the pectoral fin has been documented in a number of dolphin species, and various explanations about its function have been offered. Pectoral fin contact can take a variety of forms, and involves a number of body parts and movements, likely differing depending upon social or ecological context. For this study, we compare the pectoral fin contact behaviour of two species of wild dolphins: Indo-Pacific bottlenose dolphins (Tursiops aduncus) from around Mikura Island, Japan, and Atlantic spotted dolphins (Stenella frontalis) from The Bahamas. The two study populations exhibit surprising similarity in the ways in which pectoral fin contacts are used, despite differences in species and environmental conditions at the two sites. Differences in contact rates for calves between the two sites suggest that calf-focused aggression from adult dolphins is more prevalent at Mikura than in The Bahamas. Our results suggest that pectoral fin contact behaviour seems to be driven primarily by social pressures, and may be similar in function to allogrooming described in primates.     

Occurrence of Indo-Pacific bottlenose dolphins (Tursiops aduncus) off the Wild Coast of South Africa using photographic identification

The present study represents the first reported boat-based photographic identification study of Indo-Pacific bottlenose dolphins (Tursiops aduncus) off the Wild Coast of southeast South Africa. This area is known for the annual sardine run, which attracts apex predators to the region during the austral winter. Dedicated photo-identification surveys were conducted along this coast at three different study sites in February, June, and November of each year from 2014 to 2016. During 47 surveys, 136 bottlenose dolphin groups were encountered, an estimated 4,474 dolphins observed, and 2,149 individuals were identified. Although most individuals (N = 1,770, 82.4%) were only observed once, some were resighted 2–7 times (N = 379, 17.6%), with an average of 305 days (range: 88–705 days) between resightings. The majority of bottlenose dolphins were resighted within the same study site (N = 192), indicating some degree of residency. However, 65 individuals were observed at two different study sites, indicating individual movements along the coast. Our findings contrast earlier suggestions that bottlenose dolphins only use the Wild Coast during the sardine run, as we found large number of animals year-round with some level of site fidelity. This highlights the importance of the Wild Coast to bottlenose dolphins and provides further information on their status off southeastern South Africa.     

Spatial and Temporal Variations in the Occurrence and Foraging Activity of Coastal Dolphins in Menai Bay,Zanzibar, Tanzania

Understanding temporal patterns in distribution, occurrence and behaviour is vital for the effective conservation of cetaceans. This study used cetacean click detectors (C-PODs) to investigate spatial and temporal variation in occurrence and foraging activity of the Indo-Pacific bottlenose (Tursiops aduncus) and Indian Ocean humpback (Sousa plumbea) dolphins resident in the Menai Bay Conservation Area (MBCA), Zanzibar, Tanzania. Occurrence was measured using detection positive minutes. Inter-click intervals were used to identify terminal buzz vocalisations, allowing for analysis of foraging activity. Data were analysed in relation to spatial (location) and temporal (monsoon season, diel phase and tidal phase) variables. Results showed significantly increased occurrence and foraging activity of dolphins in southern areas and during hours of darkness. Higher occurrence at night was not explained by diel variation in echolocation rate and so were considered representative of occurrence patterns. Both tidal phase and monsoon season influenced occurrence but results varied among sites, with no general patterns found. Foraging activity was greatest during hours of darkness, High water and Flood tidal phases. Comparisons of echolocation data among sites suggested differences in the broadband click spectra of MBCA dolphins, possibly indicative of species differences. These dolphin populations are threatened by unsustainable fisheries bycatch and tourism activities. The spatial and temporal patterns identified in this study have implications for future conservation and management actions with regards to these two threats. Further, the results indicate future potential for using passive acoustics to identify and monitor the occurrence of these two species in areas where they co-exist.     

Whistling in a noisy ocean: bottlenose dolphins adjust whistle frequencies in response to real-time ambient noise levels

Common bottlenose dolphins (Tursiops truncatus) use complex acoustic behaviours for communication, group cohesion and foraging. Ambient noise from natural and anthropogenic sources has implications for the acoustic behaviour of dolphins, and research shows that average ambient noise levels alter dolphin acoustic behaviour. However, when background noise levels are highly variable, the relationships between noise and acoustic behaviour over short time periods are likely important. This study investigates whether bottlenose dolphins altered the temporal and spectral qualities of their whistles in relation to the ambient noise present at the time the whistles were produced. Dolphin groups were recorded in Tampa Bay (western Florida) between 2008 and 2015. Six whistle parameters were analysed in spectrogram software (minimum frequency, maximum frequency, bandwidth, peak frequency, duration and number of inflection points) and ambient noise levels were calculated immediately prior to each whistle. Linear regression analysis indicated that the minimum, maximum and peak frequencies of whistles had significant positive relationships with the ambient noise levels present at the time of the whistles. These models suggested that for each 1 dB increase in ambient noise, minimum frequency increased by 121 Hz, maximum frequency increased by 108 Hz and peak frequency increased by between 122 and 144 Hz. As ambient noise is typically low frequency, this suggests that bottlenose dolphins increased whistle frequency in response to real-time noise levels to avoid masking. Future research to determine the fitness consequences of noise-induced changes in the communication behaviour of dolphins would be an important contribution to conservation efforts.     

Differences in acoustic signals from Delphinids in the western North Atlantic and northern Gulf of Mexico

Whistle characteristics were quantitatively compared between both geographically separated and neighboring populations of Atlantic spotted dolphins (Stenella frontalis), bottlenose dolphins (Tursiops truncatus), and pilot whales (Globicephala spp.) in U.S. waters to evaluate if intraspecific acoustic differences exist between groups. We compared nine whistle characteristics between continental shelf and offshore Atlantic spotted dolphins in the western North Atlantic and between northern Gulf of Mexico and western North Atlantic bottlenose dolphins and pilot whales using discriminant analysis. Offshore Atlantic spotted dolphin whistles were significantly different (Hotelling's T², P= 0.0003) from continental shelf whistles in high frequency, bandwidth, duration, number of steps, and number of inflection points. Atlantic bottlenose dolphin whistles were significantly different (Hotelling's T², P < 0.0001) from those in the Gulf of Mexico in duration, number of steps, and number of inflection points. There was no significant difference between pilot whale whistles in the two basins. The whistle differences indicate acoustic divergence between groups in different areas that may arise from geographic isolation or habitat separation between neighboring but genetically distinct populations of dolphins. This study supports the premise that acoustic differences can be a tool to evaluate the ecological separation between marine mammal groups in field studies.     

Bottlenose dolphins that forage with artisanal fishermen whistle differently

Acoustic communication is a taxonomically widespread phenomenon, crucial for social animals. We evaluate social sounds from bottlenose dolphins (Tursiops truncatus) of Laguna, southern Brazil, whose social structure is organized around a cooperative foraging tactic with artisanal fishermen. This tactic involves stereotyped and coordinated behaviour by dolphins and fishermen and is performed by a subset of the dolphin population, splitting it into two distinct social communities. We compared the acoustic parameters and type of whistles emitted by dolphins of the “non‐cooperative” and “cooperative” communities, both during their interactions with fishermen and in times where dolphins were engaged in other types of foraging. Our findings show how dolphins’ social sounds differ between foraging tactics and social communities. The frequencies of six whistle types (ascending, descending, concave, convex, multiple, flat) were significantly dependent on tactics and communities. Ascending whistles were more common than expected during foraging without fishermen, and among dolphins of the non‐cooperative community. Whistle acoustic parameters (duration, number of inclination changes and inflection points, and initial, final, maximum, minimum frequencies) also varied between social communities. In general, whistles emitted by cooperative dolphins, mainly when not interacting with fishermen, tended to be shorter, had higher frequency and more inflections than those emitted by non‐cooperative dolphins. These results suggest that different whistles may convey specific information among dolphins related to foraging, which we hypothesize promote social cohesion among members of the same social community. These differences in acoustic repertoires add a new dimension of complexity to this unique human–animal interaction.     

What’s occurring? Ultrasonic signature whistle use in Welsh bottlenose dolphins (Tursiops truncatus)

Animal communication signals are diverse. The types of sounds that animals produce, and the way that information is encoded in those sounds, not only varies between species but can also vary geographically within a species. Therefore, an understanding of the vocal repertoire at the population level is important for providing insight into regional differences in vocal communication signals. One species whose vocal repertoire has received considerable attention is the bottlenose dolphin. This species is well known for its use of individually distinctive identity signals, known as signature whistles. Bottlenose dolphins use their signature whistles to broadcast their identity and to maintain contact with social companions. Signature whistles are not innate, but are learnt signals that develop within the first few months of an animal’s life. It is therefore unsurprising that studies which have characterized signature whistles in wild populations of bottlenose dolphins have provided evidence of geographic variation in signature whistle structure. Here, we describe the occurrence of signature whistles in a previously unexplored wild population of bottlenose dolphins in Cardigan Bay, Wales. We present the first occurrence of a signature whistle with an ultrasonic fundamental frequency component (>30 kHz), a frequency band that was not thought to be utilized by this species for whistle communication. We also describe the occurrence of an ultrasonic non-signature whistle. Our findings highlight the importance of conducting regional studies in order to fully quantify a species’ vocal repertoire, and call into question the efficacy of those studies that use restricted sampling rates.     

Dispersal,philopatry, and the role of fission‐fusion dynamics in bottlenose dolphins

In this quantitative study of locational and social dispersal at the individual level, we show that bottlenose dolphins (Tursiops sp.) continued to use their natal home ranges well into adulthood. Despite substantial home range overlap, mother–offspring associations decreased after weaning, particularly for sons. These data provide strong evidence for bisexual locational philopatry and mother–son avoidance in bottlenose dolphins. While bisexual locational philopatry offers the benefits of familiar social networks and foraging habitats, the costs of philopatry may be mitigated by reduced mother–offspring association, in which the risk of mother–daughter resource competition and mother–son mating is reduced. Our study highlights the advantages of high fission–fusion dynamics and longitudinal studies, and emphasizes the need for clarity when describing dispersal in this and other species.     

The structure of a bottlenose dolphin society is coupled to a unique foraging cooperation with artisanal fishermen

Diverse and localized foraging behaviours have been reported in isolated populations of many animal species around the world. In Laguna, southern Brazil, a subset of resident bottlenose dolphins (Tursiops truncatus) uses a foraging tactic involving cooperative interactions with local, beach-casting fishermen. We used individual photo-identification data to assess whether cooperative and non-cooperative dolphins were socially segregated. The social structure of the population was found to be a fission-fusion system with few non-random associations, typical for this species. However, association values were greater among cooperative dolphins than among non-cooperative dolphins or between dolphins from different foraging classes. Furthermore, the dolphin social network was divided into three modules, clustering individuals that shared or lacked the cooperative foraging tactic. Space-use patterns were not sufficient to explain this partitioning, indicating a behavioural factor. The segregation of dolphins using different foraging tactics could result from foraging behaviour driving social structure, while the closer association between dolphins engaged in the cooperation could facilitate the transmission and learning of this behavioural trait from conspecifics. This unique case of a dolphin-human interaction represents a valuable opportunity to explore hypotheses on the role of social learning in wild cetaceans.     

“Porpicide” in California: Killing of harbor porpoises (Phocoena phocoena) by coastal bottlenose dolphins (Tursiops truncatus)

Between 2007 and 2009, we witnessed three aggressive interactions between harbor porpoises and bottlenose dolphins in Monterey Bay, California. This is the first time such aggression has been documented in the Pacific, and the first time a harbor porpoise was collected immediately after witnessing its death, inflicted by bottlenose dolphins. Of the bottlenose dolphins present, 92% were males either confirmed (61%) or putative (31%). Since 2005, 44 harbor porpoise deaths inflicted by bottlenose dolphins were documented in California. Aberrant behavior was rejected as a cause of aggression, based on widespread documentation of similar behaviors in other populations of free‐ranging bottlenose dolphins. The evidence for interspecies territoriality as a form of competition for prey was weak: there is little dietary overlap and there are differences in bottlenose dolphin and harbor porpoise distribution patterns in California. Object‐oriented play was plausible as a form of practice to maintain intraspecific infanticidal skills or a form of play to maintain fighting skills between male associates. Contributing factors could be high‐testosterone levels, as attacks occurred at the height of the breeding season, and/or a skewed operational sex ratio. Ultimately, we need more information about bottlenose dolphin social structure at the time of the aggression.