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.     

Fine-scale population structure of bottlenose dolphins (Tursiops truncatus) in Tampa Bay, Florida

Some populations of coastal bottlenose dolphins ( Tursiops truncatus ) comprise discrete communities, defined by patterns of social association and long-term site fidelity. We tested the hypothesis that bottlenose dolphins in Tampa Bay, Florida, form a single community. The longitudinal study of dolphins in Sarasota Bay, adjacent to Tampa Bay, allowed us to ground-truth the definition of community and test whether our approach was robust to small sample sizes of resightings. We conducted photo-identification surveys in Tampa Bay during 1988–1993, and identified 102 dolphins with 10 or more sightings. We used hierarchical cluster analysis to examine the locations and association indices of these dolphins. We used analysis of variance (ANOVA) to test for differences in mean locations and determine whether mean coefficient of association (CoA) values within a community were higher than among communities. Dolphins in Tampa Bay clustered into five putative communities differing significantly in location and CoA values. Kernel estimates of the ranges of these five communities exhibited little overlap; some communities had no overlap at all. We conclude that five discrete communities of bottlenose dolphins exist in Tampa Bay and that such fine-scale structure may be a common feature of bottlenose dolphin populations throughout the southeastern United States.     

Site fidelity,residency, and abundance of bottlenose dolphins (Tursiops sp.) in Adelaide's coastal waters,South Australia

Little is known about the ecology and behavior of southern Australian bottlenose dolphins (Tursiops sp.). This hinders assessment of their conservation status and informed decision‐making concerning their management. We used boat‐based surveys and photo‐identification data to investigate site fidelity, residency patterns, and the abundance of southern Australian bottlenose dolphins in Adelaide's coastal waters. Sighting rates and site fidelity varied amongst individuals, and agglomerative hierarchical cluster analysis led to the categorization of individuals into one of three groups: occasional visitors, seasonal residents, or year‐round residents. Lagged identification rates indicated that these dolphins used the study area regularly from year to year following a model of emigration and reimmigration. Abundance estimates obtained from multisample closed capture‐recapture models ranged from 95 individuals (SE ± 45.20) in winter 2013 to 239 (SE ± 54.91) in summer 2014. The varying levels of site fidelity and residency, and the relatively high number of dolphins found throughout the study area highlights the Adelaide metropolitan coast as an important habitat for bottlenose dolphins. As these dolphins also appear to spend considerable time outside the study area, future research, conservation, and management efforts on this population must take into account anthropogenic activities within Adelaide's coastal waters and their adjacencies.     

Remarkably low genetic diversity and strong population structure in common bottlenose dolphins (Tursiops truncatus) from coastal waters of the Southwestern Atlantic Ocean

Knowledge about the ecology of bottlenose dolphins in the Southwestern Atlantic Ocean is scarce. Increased by-catch rates over the last decade in coastal waters of southern Brazil have raised concerns about the decline in abundance of local dolphin communities. Lack of relevant data, including information on population structure and connectivity, have hampered an assessment of the conservation status of bottlenose dolphin communities in this region. Here we combined analyses of 16 microsatellite loci and mitochondrial DNA (mtDNA) control region sequences to investigate genetic diversity, structure and connectivity in 124 biopsy samples collected over six communities of photographically identified coastal bottlenose dolphins in southern Brazil, Uruguay and central Argentina. Levels of nuclear genetic diversity were remarkably low (mean values of allelic diversity and heterozygosity across all loci were 3.6 and 0.21, respectively), a result that possibly reflects the small size of local dolphin communities. On a broad geographical scale, strong and significant genetic differentiation was found between bottlenose dolphins from southern Brazil–Uruguay (SB–U) and Bahía San Antonio (BSA), Argentina (AMOVA mtDNA Φ_ST = 0.43; nuclear F_ST = 0.46), with negligible contemporary gene flow detected based on Bayesian estimates. On a finer scale, moderate but significant differentiation (AMOVA mtDNA Φ_ST = 0.29; nuclear F_ST = 0.13) and asymmetric gene flow was detected between five neighbouring communities in SB–U. Based on the results we propose that BSA and SB–U represent two distinct evolutionarily significant units, and that communities from SB–U comprise five distinct Management Units (MUs). Under this scenario, conservation efforts should prioritize the areas in southern Brazil where dolphins from three MUs overlap in their home ranges and where by-catch rates are reportedly higher.     

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.     

Resource partitioning in a cetacean community from Uruguayan waters

The study of resource partitioning within a community is central to understanding the processes that enable interspecific competition and coexistence. This study aimed to understand the resource partitioning and habitat preferences among odontocetes in Uruguay. We analyzed stable carbon and nitrogen isotopes data from the bone tissue of common dolphins (Delphinus delphis), Fraser's dolphins (Lagenodelphis hosei), false killer whales (Pseudorca crassidens), Lahille's bottlenose dolphins (Tursiops truncatus gephyreus), Burmeister's porpoises (Phocoena spinipinnis), and franciscana dolphins (Pontoporia blainvillei). Based on the isotopic values, false killer whale individuals were divided into two different ecological groups (ecotype 1 and 2). Isotopic values also suggest that franciscana dolphins, Burmeister's porpoises, Lahille's bottlenose dolphins, and false killer whales ecotype 1 feed on the continental shelf and/or high trophic level preys. At the other end of the gradient, values suggest Fraser's dolphins and false killer whales ecotype 2 use resources from the oceanic environment and/or lower trophic preys. Isotopic niche overlap was found between the common dolphin and Lahille's bottlenose dolphin and between Lahille's bottlenose dolphin and false killer whale ecotype 1. This work provides novel basic information about the trophic habit and feeding habitat of the odontocetes inhabiting one of the most important estuaries of South America.     

MORBILLIVIRUS INFECTION IN BOTTLENOSE DOLPHINS: EVIDENCE FOR RECURRENT EPIZOOTICS IN THE WESTERN ATLANTIC AND GULF OF MEXICO

Morbillivirus infection is widespread among odontocetes of the western Atlantic and Gulf of Mexico. Serologic evidence of infection in bottlenose dolphins, Tursiops truncatus , was first detected during an epizootic along the mid-Atlantic coast in 1987. Here, we report recurrent epizootics in the coastal dolphin population since at least the early 1980s based on serological surveys and regional stranding frequencies. The first observed epizootic of this series occurred in the Indian and Banana Rivers in 1982 and was followed by others on the mid-Atlantic coast in 1987–1988 and in the Gulf of Mexico between 1992 and 1994. This temporal pattern of infection is likely facilitated by the population size and its fragmentation into relatively discrete coastal communities. Introduction of morbillivirus into a community with a sufficient number of naive hosts may precipitate an epizootic, depending on the potential for transmission within the group. Propagation of an epizootic along the coast is probably determined by frequency of contact between adjacent communities and seasonal migrations.
Morbillivirus antibodies were also detected in serum from offshore bottlenose dolphins. The sero-prevalence in the latter may be higher than in coastal dolphins because of their close association with enzootically infected pilot whales ( Globicephala spp.). Occasional contact between offshore and coastal dolphins may provide an epizootiologic link between pilot whales and coastal dolphin communities.     

Environmental and social influences on the genetic structure of bottlenose dolphins (<Emphasis Type="Italic">Tursiops aduncus</Emphasis>) in Southeastern Australia

Determining genetic connectivity of bottlenose dolphin communities helps identify evolutionary mechanisms, such as environmental and social factors, that interact to shape dispersal in highly social marine mammals. Here, we expand on a localized study that found marked genetic differentiation among resident dolphins (Tursiops aduncus) in the Port Stephens embayment and adjacent coastal communities, to include four additional communities inhabiting different environment types along the New South Wales coast, Southeastern Australia. Analysis of the mitochondrial DNA control region and seven microsatellite loci suggest the nine communities may have originated from a single ancestral population that progressively colonised the coast in a southward direction. Gene flow among communities was predominately governed by habitat type. The two enclosed embayments showed the highest level of genetic differentiation from other communities, while genetic differentiation among coastal and open embayment communities generally followed a pattern of isolation by distance. Directional bias in recent migration rates was evident, with the centrally located Hunter coast communities consisting of individuals with mixed ancestry from the Northern, Southern and Port Stephens communities. Emigration from Port Stephens was substantially higher than in the opposite direction, indicating there may be social barriers to dispersal created by Port Stephens dolphins. Our results suggest that the scale of connectivity of bottlenose dolphin communities inhabiting heterogeneous environments is likely to be affected by local habitat adaptation. This has important implications for the management of communities exposed to increasing levels of anthropogenic disturbances, such as the intensive commercial dolphin-watching industry operating in Port Stephens.     

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.     

Selection of critical habitats for bottlenose dolphins (Tursiops truncatus) based on behavioral data,in relation to marine traffic in the Istanbul Strait,Turkey

Marine traffic is a significant source of disturbance to the bottlenose dolphin population in the Istanbul Strait, Turkey. To determine the importance of this threat, behavioral data together with sighting data of both dolphins and marine vessels were assessed for 2012. The current study suggests that the Istanbul Strait is used mostly as a foraging ground for bottlenose dolphins. Nonetheless, in the same area there is intense marine traffic as well as increase of industrial fishing activities in autumn. The findings of this study indicated that high‐speed ferries and high‐speed boats were the most significant source of disturbance. Moreover, increased densities of fishing vessels resulted in a drastic decline of dolphin sightings. This study highlights that vessel type, speed, distance, and density have a cumulative negative effect on dolphins. In order to mitigate the impacts of vessels, it is necessary to establish managed areas in the Istanbul Strait. Such proposed areas should limit speed and density of marine traffic and have specific restrictions on vessel routes. We propose three different seasonal managed areas according to their values as critical habitat for bottlenose dolphins in the strait.     

Replacement dolphins? Social restructuring of a resident pod of Atlantic bottlenose dolphins,Tursiops truncatus,after two major hurricanes

Environmental variations can influence the structure of ecological communities that in turn alter the grouping and association patterns of social communities. This study compares the social structure of bottlenose dolphins in the Bahamas before and after two major hurricanes. Approximately 30% of regularly seen individuals disappeared after the hurricanes, with an equal number of immigrants arriving afterwards. The primary goal of this study was to quantitatively describe social structure changes occurring after this large‐scale emigration (or death) and subsequent immigration of individuals using the social analysis program, SOCPROG 2.3. The pre‐hurricane results revealed one community with association patterns that were consistent with previous work on this population as well as other well‐documented populations. Post‐hurricane associations revealed that the community split into two distinct units, whose members associated highly within, but rarely between units. Association patterns varied between units. Immigrants assimilated well into the population, especially males. Over half of the post‐hurricane associations involved immigrants, the majority between residents and immigrants, and primarily involving immigrant males. The costs/benefits of choosing to associate with an immigrant individual differ between males and females and may have been the driving force for the changes in social structure that occurred.     

THE SOCIAL STRUCTURE OF BOTTLENOSE DOLPHINS, TURSIOPS TRUNCATUS, IN THE BAHAMAS

The social structure of coastal ecotype bottlenose dolphins, Tursiops truncatus , is largely unknown as they inhabit regions far from shore. This study reports on a community of bottlenose dolphins ≥ 27 km from Grand Bahama Island (May-September, 1993–2002). Resident and non-resident dolphins occurred in the area. Some dolphins traveled over 320 km between communities; others showed long- term site fidelity up to 17 yr. Average group size was 3–5, and was significantly larger with calves present and significantly smaller when traveling. The half-weight index was used to determine coefficients of association (COA) for individuals of known sex annually and for pooled years. Permutation tests revealed non-random associations and presence of preferred/avoided companions in all data sets. Annual COAs were low: female-female χ= 0.31, male-male χ= 0.30, and mixed-sex χ= 0.26. Mother-calf associations showed the highest values. Some males formed strong, long-term bonds. Female COAs fluctuated with reproductive status. Using pooled data, COAs were lower and the same basic trends were evident. However, strong associations seen in the annual data were not evident in pooled data. Bottlenose dolphins that inhabit offshore, shallow water show many of the same social structure characteristics as in well-studied coastal populations.     

Connectivity in the network macrostructure of Tursiops truncatus in the Pelagos Sanctuary (NW Mediterranean Sea): does landscape matter?

The bottlenose dolphin (Tursiops truncatus Montagu, 1821) is a regularly observed species in the Mediterranean Sea, but its network organization has never been investigated on a large scale. We described the network macrostructure of the bottlenose dolphin (meta)population inhabiting the Pelagos Sanctuary (a wide protected area located in the north-western portion of the Mediterranean basin) and we analysed its connectivity in relation to the landscape traits. We pooled effort and sighting data collected by 13 different research institutions operating within the Pelagos Sanctuary from 1994 to 2011 to examine the distribution of bottlenose dolphins in the Pelagos study area and then we applied a social network analysis, investigating the association patterns of the photo-identified dolphins (806 individuals in 605 sightings). The bottlenose dolphin (meta)population inhabiting the Pelagos Sanctuary is clustered in discrete units whose borders coincide with habitat breakages. This complex structure seems to be shaped by the geo-morphological and ecological features of the landscape, through a mechanism of local specialization of the resident dolphins. Five distinct clusters were identified in the (meta)population and two of them were solid enough to be further investigated and compared. Significant differences were found in the network parameters, suggesting a different social organization of the clusters, possibly as a consequence of the different local specialization.     

RESIDENCE PATTERNS OF BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) IN THE STONO RIVER ESTUARY, CHARLESTON COUNTY, SOUTH CAROLINA, U.S.A.

Residence patterns of inshore bottlenose dolphins ( Tursiops truncatus ) in the Stono River estuary, Charleston County, South Carolina were investigated as part of a larger effort to better understand stock structure of these dolphins along the east coast of the United States. Eighty-seven small-boat surveys for bottlenose dolphins were conducted from October 1994 through January 1996. Dolphins were sighted during all surveys. Approximately 304 h were spent surveying the study area; 64% ( n = 196 h) of this time was spent observing and videotaping dolphins. A catalog, containing 112 individually identified dolphins was compiled. Thirty-two percent ( n = 36) of identified dolphins were sighted once, while 28% ( n = 31) were sighted five or more times. Nineteen percent ( n = 21) of identified dolphins were determined to be year-round residents; eight percent ( n = 9) seasonal residents. The majority (64%, n = 72) of identified dolphins were sighted in the study area during a single season or in two consecutive seasons and were classified as transients. This study documents the northernmost known site of a resident bottlenose dolphin community on the east coast of the United States, suggesting a complex bottlenose dolphin stock structure.     

Sex-specific differences in the seasonal habitat use of a coastal dolphin population

Understanding the factors that contribute to a population’s habitat use is important for conservation planners and managers to identify reasons behind a population’s distribution. Habitat use often differs between sexes, however few studies on sexually monomorphic species document this difference, resulting in misleading ecological interpretations and non-targeted management actions. The aim of this study was to test for sex-specific differences in the seasonal habitat use of Indo-Pacific bottlenose dolphins (Tursiops aduncus) off Bunbury, Australia. Systematic, boat-based, photographic identification dolphin surveys (n?=?587) were conducted across seasons over 6 years during 2007–2013. Generalised additive models explored relationships between the presence-absence of dolphins and sex, water depth and benthic habitat type. Results highlighted that: (i) habitat use differed seasonally for males and females, (ii) depth had a strong influence on habitat use, which differed between sexes for summer, winter and spring, and (iii) there were no sex differences in habitat use in autumn, which coincides with the peak breeding season. In summer and autumn dolphins were concentrated in shallow, near-shore waters predominantly over reef and sand, and in winter and spring dolphins had a broader distribution over reef and mud/silt with the use of deeper, offshore waters. This pattern is consistent with the seasonally-dependent dolphin abundance that has been documented for this population. Identification of sex differences in habitat use provides management agencies with insights to implement informed actions for the conservation of this coastal dolphin population which is forecast to decline by 50% in the next two decades.     

Mitochondrial DNA analysis of sympatric morphotypes of bottlenose dolphins (genus: Tursiops) in Chinese waters

The classification within the bottlenose dolphin (genus Tursiops) is controversial. Although many morphological variants exist, most authors have concluded that the genus is composed of a single species, Tursiops truncatus (Montagu 1821). Two distinct morphotypes of bottlenose dolphins, which have been referred to as T. truncatus and T. aduncus, exist in sympatry in Chinese waters. Comparisons of a 386-bp fragment of the mitochondrial DNA (mtDNA) control region (n = 47) indicated that the two sympatric morphotypes were genetically distinct, with seven fixed site differences and a sequence divergence of approximately 4.4%. Phylogenetic analyses using maximum likelihood, neighbour-joining and maximum parsimony approaches showed that the truncatus-type dolphins from Chinese waters were more closely related to Atlantic Ocean truncatus-type than to the sympatric aduncus-type dolphins. The Atlantic truncatus-type dolphins also shared the same diagnostic sites that separated Chinese truncatus-type from aduncus-type dolphins. The molecular data agreed completely with the morphological classifications of the specimens. This congruence is strong evidence that the sympatric morphotypes in Chinese waters are reproductively isolated and comprise two distinct species. These findings have important implications for the conservation of bottlenose dolphins in Chinese waters.     

Association patterns and population dynamics of bottlenose dolphins in the Strait of Sicily (Central Mediterranean Sea): implication for management

An understanding of the population dynamics and social organization of cetaceans is essential to manage the influence of anthropogenic activities. In this study, the population size, site fidelity and social interactions of bottlenose dolphins in the Strait of Sicily (Italy) were investigated to provide recommendations for their conservation. Mark-recapture analysis was based on the encounter histories of 103 marked dolphins from 2004 to 2015. The POPAN formulation of the Jolly–Seber model in MARK software was used to estimate the size of the super-population. Site fidelity and social organization were estimated for individuals re-sighted ≥3 times. The estimated population size was 140 (SE = 15.75; 95% CI = 106–164). Dolphins had low site fidelity, and both adults and sub-adults move outside the study area. Females with calves used the area longer than other individuals. Based on our results, dolphins’ home range likely extended beyond the study area. The mean value of the Half-Weight Association Index was low and the preferred association was by casual acquaintance. However, we found a distinct aggregation of post-parturition females during the final 2 years of the study. Therefore, the pattern of association was apparently a response to an ecological requirement, which was the possibility to breed in high productivity waters. Whether these individuals are part of a larger pelagic population is unknown; however, we can conclude that the management of only coastal waters is insufficient for the conservation of dolphins in the Strait of Sicily.     

Seasonal and Diurnal Patterns of Behavior Exhibited by Atlantic Bottlenose Dolphins (Tursiops truncatus) in the Mississippi Sound

Many of the threats to bottlenose dolphins are anthropogenic factors including overfishing, high‐speed boats, chemical runoff, and noise pollution. Having a thorough understanding of the behavior and behavioral patterns of these animals can help with conservation plans to protect this species. This study examined the behavioral states and behavioral events of Atlantic bottlenose dolphins (Tursiops truncatus) in the Mississippi Sound. The behavioral states exhibited by dolphins within the Sound were found to vary by both season and time of day. Dolphins socialized more during the spring and spent more time feeding in the fall. Feeding was highest early in the morning and decreased throughout the day, while socializing occurred at low levels in the morning and increased in the early afternoon. Two distinct forms of social behavior and multiple feeding strategies were exhibited by dolphins within the Sound. Results suggest that certain percussive behavioral events may be used to communicate motivation during transitional behavioral states. These results demonstrate the need for a more complete understanding of dolphin behavior that will facilitate the conservation of this species.