Why Do Dolphins Form Mixed‐Species Associations in the Azores?

Mixed‐species associations are temporary associations between individuals of different species that are often observed in birds, primates and cetaceans. They have been interpreted as a strategy to reduce predation risk, enhance foraging success and/or provide a social advantage. In the archipelago of the Azores, four species of dolphins are commonly involved in mixed‐species associations: the common dolphin, Delphinus delphis, the bottlenose dolphin, Tursiops truncatus, the striped dolphin, Stenella coeruleoalba, and the spotted dolphin, Stenella frontalis. In order to understand the reasons why dolphins associate, we analysed field data collected since 1999 by research scientists and trained observers placed onboard fishing vessels. In total, 113 mixed‐species groups were observed out of 5720 sightings. The temporal distribution, habitat (water depth, distance to the coast), behaviour (i.e. feeding, travelling, socializing), size and composition of mixed‐species groups were compared with those of single‐species groups. Results did not support the predation avoidance hypothesis and gave little support to the social advantage hypothesis. The foraging advantage hypothesis was the most convincing. However, the benefits of mixed‐species associations appeared to depend on the species. Associations were likely to be opportunistic in the larger bottlenose dolphin, while there seemed to be some evolutionary constraints favouring associations in the rarer striped dolphin. Comparison with previous studies suggests that the formation of mixed‐species groups depends on several environmental factors, and therefore may constitute an adaptive response.     

Fine‐scale habitat use in Indo‐Pacific humpback dolphins,Sousa chinensis,may be more influenced by fish rather than vessels in the Pearl River Estuary,China

Shifts in habitat use and distribution patterns in dolphins are often concerns that can result from habitat degradation. We investigated how potential changes to a habitat from human activity may alter dolphin distributions within Lingding Bay in the Pearl River Estuary, China, by studying the relationship between fish choruses, vessel presence and Indo‐Pacific humpback dolphin (Sousa chinensis) detection rates. Analyses revealed temporal and spatial variation within fish choruses, vessel presence and dolphin detection rates. After accounting for any temporal autocorrelation, correlations between fish choruses and dolphin detection rates were also found; however, no relationship between fish choruses and vessel presence or dolphin detection rates and vessel presence were observed. Furthermore, fewer dolphins were detected at sites where fish activity was less intense. Thus fish activity, rather than vessels, may be a key factor influencing the distribution of the dolphins within the estuary. These findings emphasize the risk of potential shifts in habitat use for Indo‐Pacific humpback dolphins due to detrimental changes to prey availability and dolphin feeding grounds from human activity, such as overfishing and coastal developments, within the estuary. This is a critical conservation issue for this dolphin population that is facing intense anthropogenic pressure.     

Environmental Niche Overlap between Common and Dusky Dolphins in North Patagonia,Argentina

Research on the ecology of sympatric dolphins has increased worldwide in recent decades. However, many dolphin associations such as that between common dolphins (Delphinus delphis) and dusky dolphins (Lagenorhynchus obscurus) are poorly understood. The present study was conducted in the San Matías Gulf (SMG) ecosystem (North Patagonia, Argentina) where a high diet overlap among both species was found. The main objective of the present work was to explore the niche overlap of common and dusky dolphins in the habitat and temporal dimensions. The specific aims were (a) to evaluate the habitat use strategies of both species through a comparison of their group attributes (social composition, size and activity), and (b) to evaluate their habitat preferences and habitat overlap through Environmental Niche modeling considering two oceanographic seasons. To accomplish these aims, we used a historic database of opportunistic and systematic records collected from 1983 to 2011. Common and dusky dolphins exhibited similar patterns of group size (from less than 10 to more than 100 individuals), activity (both species use the area to feed, nurse, and copulate), and composition (adults, juveniles, and mothers with calves were observed for both species). Also, both species were observed travelling and feeding in mixed-species groups. Specific overlap indices were higher for common dolphins than for dusky dolphins, but all indices were low, suggesting that they are mainly segregated in the habitat dimension. In the case of common dolphins, the best habitats were located in the northwest of the gulf far from the coast. In the warm season they prefer areas with temperate sea surface and in the cold season they prefer areas with relatively high variability of sea surface temperature. Meanwhile, dusky dolphins prefer areas with steep slopes close to the coast in the southwestern sector of the gulf in both seasons.     

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.     

Population status and habitat occupancy of endangered river dolphins in the Karnali River system of Nepal during low water season

Ganges river dolphin abundance has undergone a predominant decline across its range since monitoring began. In Nepal, disappearance from some of the rivers it once used has already occurred. Today this species can only be found in three river systems in Nepal, the Karnali, Sapta Koshi, and Narayani, but numbers are low in these locations. To determine the abundance of dolphins remaining in the Karnali system (which includes the Karnali, Geruwa, and Mohana), and factors affecting dolphin habitat use, we conducted surveys where we recorded dolphin presence. Dolphins within this river system were sighted only in the Karnali and an abundance estimate of 5.04 ± 0.753 SE was calculated. This pattern of ranging differed from that previously reported (from previous sightings only in the Geruwa to current sightings only in the Karnali). River depth likely contributed to the presence or absence of dolphins. Shifts in available habitat between the Geruwa and Karnali have resulted from changes in the course of the main stream Karnali following construction of the Chisapani irrigation intake. Because of the low numbers of dolphins reported, there is great concern that loss of this species in Nepal is likely in the near future.     

Human Interactions with Free-Ranging and Captive Bottlenose Dolphins

ABSTRACT

The popular demand for encounters with dolphins has resulted in organized encounters around the world involving in-water interactions with both captive and free-ranging dolphins. Many concerns about these interactions have been raised with regard to the well-being of human and dolphin participants (e.g. Capaldo 1989; Iannuzzi and Rowan 1991). In this paper we (a) review existing information on human/dolphin interactions, (b) provide a qualitative comparison of such interactions in the captive and in the wild with the same species of dolphin (Tursiops truncatus), and (c) make suggestions regarding future research needs regarding the wellbeing of dolphins and humans in interactive programs.

This study found that the free-ranging dolphin/human interactions consisted primarily of basic elements of intraspecific interactions among free-ranging dolphins. Interactive dolphin behaviors in the captive environment consisted primarily of actions that had been signaled and then reinforced by humans. Potential indicators of disturbance in the dolphins were observed in both contexts, but most frequently in the captive environment. We conclude that future research is needed to assess the risks to human and dolphin participants during in-water encounters.     

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.     

SHARK ATTACKS ON BOTTLENOSE DOLPHINS (TURSIOPS ADUNCUS) IN SHARK BAY, WESTERN AUSTRALIA: ATTACK RATE, BITE SCAR FREQUENCIES, AND ATTACK SEASONALITY

Shark predation may have been a central factor influencing the evolution of sociality in dolphins, as well as a determinant of dolphin habitat use and behavior. To understand the role of predation in driving interpopulation differences in behavior and sociality, it is important to quantify differences in predation risk among populations. This study describes the frequency of shark-inflicted scars and estimates the shark attack rate on bottlenose dolphins ( Tursiops aduncus ) in Shark Bay, Western Australia. Shark bite scars were found on 74.2% (95 of 128) of non-calves, and most of these scars were inflicted by tiger sharks ( Galeocerdo cuvier ). Although there were no differences among age/sex classes in the frequency of scarring, significantly more adult males than adult females bore multiple scars. The rate of unsuccessful shark attack was estimated to be between 11% and 13% of dolphins attacked each year. Large sharks (>3 m) were responsible for a disproportionate number of attacks. However, bites from small carcharhinid sharks on 6.2% of dolphins suggest that some of these small sharks may be dolphin ectoparasites. Both the scar frequencies and attack rate suggest that Shark Bay dolphins face a greater risk of predation than bottlenose dolphins in other locations.     

Predictive Modeling of Spinner Dolphin (Stenella longirostris) Resting Habitat in the Main Hawaiian Islands

Predictive habitat models can provide critical information that is necessary in many conservation applications. Using Maximum Entropy modeling, we characterized habitat relationships and generated spatial predictions of spinner dolphin (Stenella longirostris) resting habitat in the main Hawaiian Islands. Spinner dolphins in Hawai'i exhibit predictable daily movements, using inshore bays as resting habitat during daylight hours and foraging in offshore waters at night. There are growing concerns regarding the effects of human activities on spinner dolphins resting in coastal areas. However, the environmental factors that define suitable resting habitat remain unclear and must be assessed and quantified in order to properly address interactions between humans and spinner dolphins. We used a series of dolphin sightings from recent surveys in the main Hawaiian Islands and a suite of environmental variables hypothesized as being important to resting habitat to model spinner dolphin resting habitat. The model performed well in predicting resting habitat and indicated that proximity to deep water foraging areas, depth, the proportion of bays with shallow depths, and rugosity were important predictors of spinner dolphin habitat. Predicted locations of suitable spinner dolphin resting habitat provided in this study indicate areas where future survey efforts should be focused and highlight potential areas of conflict with human activities. This study provides an example of a presence-only habitat model used to inform the management of a species for which patterns of habitat availability are poorly understood.     

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.     

The use of auditory stimulants during swim encounters with Hector's dolphins (Cephalorhynchus hectori hectori) in Akaroa Harbour,New Zealand

The coastal distribution of Hector's dolphins and their attraction to vessels make them easily accessible to commercial tour operations. For over 25 yr, tour operators have been undertaking view and swim‐with‐dolphin trips in Akaroa Harbour, New Zealand. Since 2003, auditory stimulants, in particular stones, have been provided during such swim encounters. The potential effects associated with such stimulants have not, until now, been examined. Here, we investigate the effects of stones and other human‐induced noise on Hector's dolphin behavior. The use of stones significantly affected how dolphins interacted with swimmers. Specifically, swimmers who used stones had a greater probability of close approaches by dolphins than those who sang or simply floated on the surface of the water. The number of close and sustained approaches was also significantly higher for swimmers using stones. Dolphins were more interactive with active swimmers, approaching closer and engaging for longer than with nonactive swimmers. Dolphins socializing had a tendency to be engaged longer with swimmers. The use of stones as an auditory stimulant to sustain or enhance interactions with dolphins by artificial means may not be in the best interest of an endangered species, which already faces a range of challenges due to human activity.     

Does tiger shark predation risk influence foraging habitat use by bottlenose dolphins at multiple spatial scales?

Prey availability and predation risk are important determinants of habitat use, but their importance may vary across spatial scales. In many marine systems, predator and prey distributions covary at large spatial scales, but do no coincide at small spatial scales. We investigated the influences of prey abundance and tiger shark ( Galeocerdo cuvier ) predation risk on Indian Ocean bottlenose dolphin ( Tursiops aduncus ) habitat use across multiple spatial scales, in Shark Bay, Western Australia. Dolphins were distributed between deep and shallow habitats and across microhabitats within patches approximately proportional to prey density when shark abundance was low. When shark abundance was high, foraging dolphins greatly reduced their use of dangerous, but productive, shallow patches relative to safer deep ones. Also, dolphins reduced their use of interior portions of shallow patches relative to their edges, which have higher predator density but lower intrinsic risk (i.e. a higher probability of escape in an encounter situation). These results suggest that predation risk and prey availability influence dolphin habitat use at multiple spatial scales, but intrinsic habitat risk, and not just predator encounter rate, is important in shaping dolphin space use decisions. Therefore, studies of habitat use at multiple spatial scales can benefit from integrating data on prey availability and the subcomponents of predation risk.     

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.     

Behavioural Effects of Tourism on Oceanic Common Dolphins,Delphinus sp., in New Zealand: The Effects of Markov Analysis Variations and Current Tour Operator Compliance with Regulations

Common dolphins, Delphinus sp., are one of the marine mammal species tourism operations in New Zealand focus on. While effects of cetacean-watching activities have previously been examined in coastal regions in New Zealand, this study is the first to investigate effects of commercial tourism and recreational vessels on common dolphins in an open oceanic habitat. Observations from both an independent research vessel and aboard commercial tour vessels operating off the central and east coast Bay of Plenty, North Island, New Zealand were used to assess dolphin behaviour and record the level of compliance by permitted commercial tour operators and private recreational vessels with New Zealand regulations. Dolphin behaviour was assessed using two different approaches to Markov chain analysis in order to examine variation of responses of dolphins to vessels. Results showed that, regardless of the variance in Markov methods, dolphin foraging behaviour was significantly altered by boat interactions. Dolphins spent less time foraging during interactions and took significantly longer to return to foraging once disrupted by vessel presence. This research raises concerns about the potential disruption to feeding, a biologically critical behaviour. This may be particularly important in an open oceanic habitat, where prey resources are typically widely dispersed and unpredictable in abundance. Furthermore, because tourism in this region focuses on common dolphins transiting between adjacent coastal locations, the potential for cumulative effects could exacerbate the local effects demonstrated in this study. While the overall level of compliance by commercial operators was relatively high, non-compliance to the regulations was observed with time restriction, number or speed of vessels interacting with dolphins not being respected. Additionally, prohibited swimming with calves did occur. The effects shown in this study should be carefully considered within conservation management plans, in order to reduce the risk of detrimental effects on common dolphins within the region.     

Associations between seabirds and cetaceans: a review

(1) Seabird-cetacean associations involve many genera.
(2) Some seabirds/cetaceans associate with more species than others, e.g. minke and pilot whales, Common dolphins and porpoises amongst cetaceans, and gannet & kittiwake amongst seabirds. Killer whale, sperm whale, shag and cormorant have not been recorded in associations, in the N.E. Atlantic.
(3) All observed associations would be predicted on the basis of the diets of the associated species.
(4) Most associations are probably opportunistic or incidental, as a result of concentration of shared prey.
(5) Some species, however, may associate more regularly, e.g. minke & pilot whales, Common dolphin and porpoise amongst cetaceans and gannet amongst seabirds. It is not possible to say which derives benefit from the association, but on theoretical grounds it is most likely to be the seabird.
(6) Minke and pilot whales may drive towards the surface food normally out of reach of seabirds, particularly Procellariformes. Common dolphins feed by herding fast-moving fish shoals into tight groups which will be a more concentrated food source for diving sea-birds, such as gannets, and make food more accessible to other species. Porpoises may achieve the same result with inshore shoals of sprats and sand eels, particularly for auks and kittiwakes, and terns.     

Assessing spatial patterns and density of a dolphin population through signature whistles

Some dolphin species produce signature whistles, which may allow the identification of individual dolphins using passive acoustic monitoring (PAM). Identifying individuals by their sounds may enhance the opportunities for monitoring and addressing biological and ecological questions about these species. Here, we explored the potential of signature whistles to investigate ecological aspects of a resident bottlenose dolphin population. Using a limited data set, with few individuals recognized by signature whistles, combined with spatial capture-recapture (SCR) methods, we investigated how effective such approach is describing spatial use patterns and estimating density for this population. The data were collected using 4–6 stationary bottom-moored recorders. Since only eight signature whistles were identified, our density estimate may represent a subset of the entire population. However, even with only a few signature whistles identified, our results confirmed the center of the core area used by these dolphins as the area with the highest encounter probability. In addition, our results provided evidence that these dolphins have the same spatial use pattern at night as during the day. This study shows that SCR analysis of signature whistle data can improve our ecological knowledge and understanding of dolphin populations.     

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.     

Abundance and residency dynamics of the Indo-Pacific humpback dolphin,Sousa chinensis,in the Dafengjiang River Estuary,China

Robust population size estimates are essential for informing population conservation status. Residency dynamics show population habitat use through time. Population size of Indo-Pacific humpback dolphins (Sousa chinensis) has been extensively investigated in Chinese waters, but their residency dynamics are rarely known. Mark-recapture analysis based on photo-identification records was applied to humpback dolphins in the Dafengjiang River Estuary habitat, one of the key habitats in the northern Beibu Gulf, China. Movement analyses based on lagged identification rate indicated the humpback dolphins spent, on average, 78.5 days inside and 46.9 days outside the survey area. Within the study area, the humpback dolphin abundance was 83 identifiable dolphins. A total of 353–430 humpback dolphins, estimated by POPAN modeling, were involved in this fluid habitat-use dynamic. Robust Design analysis showed strong seasonality in humpback dolphin abundance and emigration probability, implying a movement- and habitat-use pattern likely associated with spatiotemporal distribution of oceanographic characteristics and prey occurrences. Population surveys and conservation measures currently conducted in Chinese waters seldom consider seasonality in movements between habitat patches, which can be addressed by genetic analyses across habitats and cross-matching photo-identification records among neighboring habitats.