Evidence for infanticide in bottlenose dolphins: an explanation for violent interactions with harbour porpoises?

Most harbour porpoises found dead on the north-east coast of Scotland show signs of attack by sympatric bottlenose dolphins, but the reason(s) for these violent interactions remain(s) unclear. Post-mortem examinations of stranded bottlenose dolphins indicate that five out of eight young calves from this same area were also killed by bottlenose dolphins. These data, together with direct observations of an aggressive interaction between an adult bottlenose dolphin and a dead bottlenose dolphin calf, provide strong evidence for infanticide in this population. The similarity in the size range of harbour porpoises and dolphin calves that showed signs of attack by bottlenose dolphins suggests that previously reported interspecific interactions could be related to this infanticidal behaviour. These findings appear to provide the first evidence of infanticide in cetaceans (whales, dolphins and porpoises). We suggest that infanticide must be considered as a factor shaping sociality in this and other species of cetaceans, and may have serious consequences for the viability of small populations.     

ECOLOGY, BEHAVIOR AND SOCIAL ORGANIZATION OF THE BOTTLENOSE DOLPHIN: A REVIEW

The authors review the literature on bottlenose dolphin ecology, behavior and social organization, focusing on data collected on free-ranging animals. Most bottlenose dolphins studied to date have had definable home ranges, and behavioral, morphological and biochemical information indicates discrete stocks in some areas. Bottlenose dolphins appear to form relatively permanent social groups based on sex and age. Mother—calf bonds are long-lasting. Movement patterns are extremely variable from location to location but are relatively predictable at any given site. Food resources are one of the most important factors affecting movements. Bottlenose dolphin behavior is very flexible, and these dolphins are generally active day and night. Feeding peaks in the morning and afternoon have been observed at several sites. Social behavior is an important component of daily activities. Sharks are the most significant predator on bottlenose dolphins in most areas, but captive and wild studies show that dolphins and sharks frequently live in harmony as well. Human activities may be helpful, harmful or neutral to bottlenose dolphins, but interactions with humans are frequent for these coastal cetaceans.     

SHORT-TERM EFFECTS OF BOAT TRAFFIC ON BOTTLENOSE DOLPHINS, TURSIOPS TRUNCATUS, IN SARASOTA BAY, FLORIDA

Coastal cetaceans are subject to potential injury or disturbance from vessels. In Sarasota, Florida, where about 120 resident bottlenose dolphins, Tursiops truncatus , share the inshore waters with over 34,000 registered boats, disturbance potential is high. We assessed specific behavioral responses of individual dolphins to boat traffic. We conducted focal animal behavioral observations during opportunistic and experimental boat approaches involving 33 well-known identifiable individual bottlenose dolphins. Dolphins had longer interbreath intervals (IBI) during boat approaches compared to control periods (no boats within 100 m). Treatment IBI length was inversely correlated with distance to the nearest boat in opportunistic observations. During 58 experimental approaches to 18 individuals, a video system suspended from a tethered airship was used to observe subsurface responses of focal dolphins as boats under our control, operating at specified speeds, were directed near dolphins. Dolphins decreased interanimal distance, changed heading, and increased swimming speed significantly more often in response to an approaching vessel than during control periods. Probability of change for both interanimal distance and heading increased when dolphins were approached while in shallow water. Our findings provide additional support for the need to consider disturbance in management plans for cetacean conservation.     

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.     

CETACEANS OF THE NORTHERN GULF OF CALIFORNIA: DISTRIBUTION, OCCURRENCE, and RELATIVE ABUNDANCE

A total of 1,715 km of boat-based surveys and 1,521 km of aircraft-based surveys was conducted from 1986–1989 to assess the distribution, relative abundance, and ecological relationships of cetaceans in the northern Gulf of California. Seven cetacean species were seen; in decreasing frequency of groups encountered they were: bottlenose dolphins, Tursiops truncatus ; vaquitas, Phocoena sinus ; common dolphins, Delphinus delphis ; fin whales, Balaenoptera physalus ; Bryde's whales, B. edeni ; gray whales, Eschrichtius robustus , and killer whales, Orcinus orca. Common dolphins were numerically dominant and bottlenose dolphins were seen most often. Bryde's whales and vaquitas had the smallest group sizes. In general, the odontocete cetaceans were separated spatially, whereas the distribution of Bryde's and fin whales overlapped considerably.     

Parallel influence of climate on the behaviour of Pacific killer whales and Atlantic bottlenose dolphins

The grouping behaviour of animals is governed by intrinsic and extrinsic factors which play an important role in shaping their social organization. We investigated the influence of ocean climate variation on the grouping behaviour of two widely separated populations of cetaceans, inhabiting north Atlantic and north Pacific coastal waters. The group size of both bottlenose dolphins in the Moray Firth, UK, and killer whales in Johnstone Strait, Canada, varied from year to year in relation to large‐scale ocean climate variation. Local indices of prey abundance were also related both to climate indices and predator group sizes. The cetaceans tended to live in smaller groups when there was less salmon available in both areas which seem to occur 2 years after a lower phase of the North Atlantic and Pacific Decadal Oscillations. These findings suggest that, even in highly social mammals, climate variation may influence social organization through changes in prey availability.     

Dolphin genome provides evidence for adaptive evolution of nervous system genes and a molecular rate slowdown

Cetaceans (dolphins and whales) have undergone a radical transformation from the original mammalian bodyplan. In addition, some cetaceans have evolved large brains and complex cognitive capacities. We compared approximately 10 000 protein-coding genes culled from the bottlenose dolphin genome with nine other genomes to reveal molecular correlates of the remarkable phenotypic features of these aquatic mammals. Evolutionary analyses demonstrated that the overall synonymous substitution rate in dolphins has slowed compared with other studied mammals, and is within the range of primates and elephants. We also discovered 228 genes potentially under positive selection (dN/dS > 1) in the dolphin lineage. Twenty-seven of these genes are associated with the nervous system, including those related to human intellectual disabilities, synaptic plasticity and sleep. In addition, genes expressed in the mitochondrion have a significantly higher mean dN/dS ratio in the dolphin lineage than others examined, indicating evolution in energy metabolism. We encountered selection in other genes potentially related to cetacean adaptations such as glucose and lipid metabolism, dermal and lung development, and the cardiovascular system. This study underlines the parallel molecular trajectory of cetaceans with other mammalian groups possessing large brains.     

Historical and current records of aquarium cetaceans in China

The number of cetaceans housed in aquariums in China is increasing. Detailed information on the historical and current population status has not been reported, despite its importance for successful breeding and population management. Questionnaires were conducted between December 2006 and May 2009, and the information was used to construct studbooks. Our survey showed that 10 species had been introduced to aquariums since 1978, including 26 (with 15 in the current population) finless porpoises (Neophocaena phocaenoides), 5 (5) false killer whales (Pseudorca crassidens), 94 (80) common bottlenose dolphins (Tursiops truncatus), 48 (30) Indo-Pacific bottlenose dolphins (Tursiops aduncus), 36 (32) beluga whales (Delphinapterus leucas), 10 (10) pantropical spotted dolphins (Stenella attenuata), 8 (8) Risso's dolphins (Grampus griseus), 2 (2) short-finned pilot whales (Globicephala macrorhynchus), 2 (2) Pacific white-sided dolphins (Lagenorhynchus obliquidens), and 5 (0) baiji dolphins (Lipotes vexillifer). The number of cetaceans has increased markedly in the past 32 years, especially since 1995. Currently, 184 individuals are under human care throughout China, a number larger than any other country with an International Species Information System membership. In addition, the Annual Survival Rates of bottlenose dolphins (0.959) and beluga whales (0.968) were found higher than those reported previously (0.93-0.951 and 0.94-0.954, respectively).     

Molecular cloning and histological localization of LH-like substances in a bottlenose dolphin (Tursiops truncatus) placenta

All mammals exhibit pituitary-specific expression of LH and FSH, whereas placental expression of gonadotropins has been reported only in primates and equids. Some cetaceans, such as dolphins, have a long gestational period and a sexual cycle of about 27 days almost comparable with that of humans. Histologically, dolphins have an epitheliochorial placentae that resembles placentas of Perissodactyla including horses. In the present study, we cloned cDNAs encoding gonadotropins and observed their immunohistochemical localization in the placenta of bottlenose dolphin. The cDNAs obtained encoded 120 amino acids for the alpha-subunit (including 96 amino acids of mature proteins), and 141 amino acids for the beta-subunit (including 121 amino acids of mature proteins). The sequence of the alpha-subunit was similar to that in the pig (Artiodactyla) pituitary glycoprotein hormone [96.7% homology at amino acids (aa) level], and the sequence of the beta-subunit was similar to that of luteinizing hormone (LH) in the pig [94.3% homology at aa level] and white rhinoceros (Perissodactyla) [93.3% homology at aa level]. Of interest, dolphin LHbeta lacks carboxyl-terminal-peptides (CTP). This fact suggests that CTP are not essential for placental expression of gonadotropin in dolphins. Immunohistochemical observations employing anti-ovine LHbeta antibody revealed positive staining in the villositycal tissue. Our observations suggest placental expression of gonadotropin homologues in cetaceans and possible evolutionary conservation of placentae-derived hormonal control of ovarian functions during pregnancy.     

Ecology and conservation of common bottlenose dolphins Tursiops truncatus in the Mediterranean Sea

• 1 Bottlenose dolphins Tursiops truncatus are amongst the best‐known cetaceans. In the Mediterranean Sea, however, modern field studies of cetaceans did not start until the late 1980s. Bottlenose dolphins have been studied only in relatively small portions of the basin, and wide areas remain largely unexplored.
• 2 This paper reviews the ecology, behaviour, interactions with fisheries and conservation status of Mediterranean bottlenose dolphins, and identifies threats likely to have affected them in historical and recent times.
• 3 Whilst intentional killing was probably the most important cause of mortality until the 1960s, important ongoing threats include incidental mortality in fishing gear and the reduced availability of key prey caused by overfishing and environmental degradation throughout the region. Additional potential or likely threats include the toxic effects of xenobiotic chemicals, epizootic outbreaks, direct disturbance from boating and shipping, noise, and the consequences of climate change.
• 4 The flexible social organization and opportunistic diet and behaviour of bottlenose dolphins may allow them to withstand at least some of the effects of overfishing and habitat degradation. However, dolphin abundance is thought to have declined considerably in the region and management measures are needed to prevent further decline.
• 5 Management strategies that could benefit bottlenose dolphins, such as sustainable fishing, curbing marine pollution and protecting biodiversity, are already embedded in legislation and treaties. Compliance with those existing commitments and obligations should be given high priority.

     

Stable isotopes differentiate bottlenose dolphins off west-central Florida

Distinguishing discrete population units among continuously distributed coastal small cetaceans is challenging and crucial to conservation. We evaluated the utility of stable isotopes in assessing group membership in bottlenose dolphins (Tursiops truncatus) off west-central Florida by analyzing carbon, nitrogen, and sulfur isotope values (δ¹³C, δ¹⁵N, and δ³⁴S) of tooth collagen from stranded dolphins. Individuals derived from three putative general population units: Sarasota Bay (SB), nearshore Gulf of Mexico (GULF), and offshore waters (OFF). Animals of known history (SB) served to ground truth the approach against animals of unknown history from the Gulf of Mexico (GULF, OFF). Dolphin groups differed significantly for each isotope. Average δ¹³C values from SB dolphins (−10.6‰) utilizing sea grass ecosystems differed from those of GULF (−11.9‰) and OFF (−11.9‰). Average δ¹⁵N values of GULF (12.7‰) and OFF (13.2‰) were higher than those of SB dolphins (11.9‰), consistent with differences in prey trophic levels. δ³⁴S values showed definitive differences among SB (7.1‰), GULF (11.3‰), and OFF (16.5‰) dolphins. This is the first application of isotopes to population assignment of bottlenose dolphins in the Gulf of Mexico and results suggest that isotopes may provide a powerful tool in the conservation of small cetaceans.     

High gene flow in oceanic bottlenose dolphins (Tursiops truncatus) of the North Atlantic

Despite the openness of the oceanic environment, limited dispersal and tight social structure often induce genetic structuring in marine organisms, even in large animals such as cetaceans. In the bottlenose dolphin, mitochondrial and nuclear DNA analyses have revealed the existence of genetic differentiation between pelagic (or offshore) and coastal (or nearshore) ecotypes in the western North Atlantic, as well as between coastal populations. Because previous studies concentrated on continental margins, we analysed the population structure of bottlenose dolphins in two of the most isolated archipelagos of the North Atlantic: the Azores and Madeira. We analysed 112 samples collected on live animals in the two archipelagos, and nine samples collected on stranded animals in Madeira and mainland Portugal. Genetic analyses consisted in molecular sexing, sequencing of part of the mitochondrial hyper-variable region, and screening of ten microsatellite loci. We predicted that: (1) there is at least one pelagic and one or more coastal populations in each archipelago; (2) populations are differentiated between and possibly within archipelagos. Contrary to these predictions, results indicated a lack of population structure in the study area. In addition, comparison with published sequences revealed that the samples from the Azores and Madeira were not significantly differentiated from samples of the pelagic population of the western North Atlantic. Thus, bottlenose dolphins occurring in the pelagic waters of the North Atlantic belong to a large oceanic population, which should be regarded as a single conservation unit. Unlike what is known for coastal populations, oceanic bottlenose dolphins are able to maintain high levels of gene flow.     

Genetic evidence for sex-biased dispersal in resident bottlenose dolphins (Tursiops aduncus)

In most mammals males usually disperse before breeding, while females remain in their natal group or area. However, in odontocete cetaceans behavioural and/or genetic evidence from populations of four species indicate that both males and females remain in their natal group or site. For coastal resident bottlenose dolphins field data suggest that both sexes are philopatric to their natal site. Assignment tests and analyses of relatedness based on microsatellite markers were used to investigate this hypothesis in resident bottlenose dolphins, Tursiops aduncus, from two small coastal populations of southeastern Australia. Mean corrected assignment and mean relatedness were higher for resident females than for resident males. Only 8% of resident females had a lower probability than average of being born locally compared to 33% of resident males. Our genetic data contradict the hypothesis of bisexual philopatry to natal site and suggest that these bottlenose dolphins are not unusual amongst mammals, with females being the more philopatric and males the more dispersing sex.     

Post-mortem investigations on stranded dolphins and porpoises from Hong Kong waters

Stranded cetaceans reported from the territorial waters of Hong Kong during the period May 1993 to March 1998 were examined to establish factors that may have contributed to their death. During the current study, 28 Indo-Pacific hump-backed dolphins (Sousa chinensis), 32 finless porpoises (Neophocaena phocaenoides), and four bottlenose dolphins (Tursiops truncatus) were necropsied. Bacteria (15 species) were isolated from nine animals. Of these bacteria, 47% were of possible fecal origin reflecting the high level of sewage contamination in Hong Kong's waters. One finless porpoise displayed wounds caused by a shark attack, and two female finless porpoises presented prolapsed uteri. At least 10 finless porpoises showed evidence of moderate to heavy lungworm infections (Halocercus pingi), and this appears to have been a factor contributing to death in at least six animals. Evidence suggesting blunt traumatic injury (probably caused by boat collisions) was found in six cetaceans (three finless porpoises and three hump-backed dolphins). Signs of fishery-related mortality were detected in at least nine animals (six hump-backed dolphins, two finless porpoises, and one bottlenose dolphin). Of these two human-caused mortality types, pre-existing disease or bacterial infection were detected in 29% of cases. Results indicate that human factors may have played a significant role in the death of at least 15 animals (32% of hump-backed dolphins, 15% of finless porpoises, and 25% of bottlenose dolphins).     

Citrobacter freundii septicemia in a stranded newborn Cuvier's beaked whale (Ziphius cavirostris)

Citrobacter freundii, a gram-negative enterobacterium, may cause fatal septicemia in humans and animals. Its potential pathogenic role in cetaceans (bottlenose dolphins and beluga whales) has been hypothesized. Here we describe fatal C. freundii septicemia in a stranded newborn Cuvier's beaked whale (Ziphius cavirostris).     

Quantitative Comparison of Whistle Repertoires from Captive Adult Bottlenose Dolphins (Delphinidae,Tursiops truncatus): a Re-evaluation of the Signature Whistle Hypothesis

The prevailing view among researchers of dolphin communication is that bottlenose dolphins possess an individualized whistle contour; known as the ‘signature whistle’, it accounts for 74–95 % of a dolphin's whistle repertoire and functions to signal the individual identity of the whistler. This study used a new quantitative technique, termed the contour similarity technique (CS technique), and reports on the quantitative comparison of whistles from the individuals of three different social groups of bottlenose dolphins in socially interactive contexts. Results suggest that captive adult dolphins share several different whistle types including one predominant whistle type shared by all individuals across three different social groups. These analyses suggest a different interpretation of the dolphin whistle repertoire than has previously been proposed by proponents of the signature whistle hypothesis. In addition, results from our study support the results of early studies, published before the advent of the signature whistle hypothesis, in which investigators reported a large whistle repertoire within socially interactive captive and free-ranging groups and a predominant whistle type similar to that found in our study. Our results, combined with the results from earlier studies of dolphin vocal behaviour, suggest that the signature whistle hypothesis is incomplete and that dolphin whistle repertoires need to be analysed with respect to behavioural context and social relationships. In addition, these results suggest that contour discrimination and other acoustic features of whistles need to be tested in perception and categorization experiments.     

Struvite calculus in the vagina of a bottlenose dolphin (Tursiops truncatus)

On 27 January 2000, a struvite calculus was observed in the vagina during necropsy of a 138-cm-long female bottlenose dolphin (Tursiops truncatus) collected from the Stono River, South Carolina (USA). Vaginal calculi have been reported in other species of cetaceans but not in bottlenose dolphins. Urinary tract infection might have been an underlying cause of the calculus. While urinary tract inflammation was not detected by light microscopic evaluation of sections of the urinary tract, it is conceivable that sufficient time had lapsed following voiding of the calculus through the urethra for urinary tract infection to have resolved. To further define the prevalence and significance of urolithiasis, prosectors of dead stranded marine mammals are encouraged to closely observe their urinary and genital tracts for calculi and to submit them for quantitative analysis.     

Sighting frequency and relative abundance of bottlenose dolphins (Tursiops truncatus) along the northeast coast of Margarita Island and Los Frailes Archipelago, Venezuela

The study of local cetaceans in Venezuela has a very recent history, and few efforts have been made in the assessment of coastal populations based on field research. The occurrence of whales and dolphins along the northeast coast of Venezuela has been documented through sightings and stranding records. Given the underwater topographical features and the influence of upwelling processes, this area is considered a very productive coastal ecosystem. Our objective was to establish the sighting frequency and relative abundance of bottlenose dolphins in the area. Sighting records were gathered on bottlenose dolphins and other cetacean species occurring along the northeast coast of Margarita Island and Los Frailes Archipelago through direct observation during land-based (6 surveys, 48 hours of observation) and boat-based surveys (24 surveys, 121 hours of observation, 1295 km covered). A sighting frequency was calculated using two methodologies and then compared, considering: 1) a mean effective observation time (4.27 hours), and 2) distance covered with cetacean sightings (1108 kin). A third method is proposed relating a mean effective distance covered with cetacean sightings and expressed as a percentage. The abundance index was calculated using the mean effective observation time. The sighting frequency of Tursiops truncattus in the study area was 3 - 4 sightings per day of 4.27 observation hours, or by 185 kilometers covered. The relative abundance was calculated as 35 dolphins in the study area, so a total population of less than 60 dolphins could inhabit the proposed range. Tursiops truncatus is the dominant species in the northeast coast of Margarita Island and Los Frailes Archipelago with 70% of all the sightings, so this locality could be termed as the distribution range of a possible local population of bottlenose dolphins.     

台湾海峡厦门-东山水域瓶鼻海豚的种群密度、分布和误捕研究

台湾海峡厦门东山水域瓶鼻海豚 ( Tursiopsaduncus)的种群密度约为 0 .0 4 36± 0 .0 2 86头 /km2 ,该水域南部瓶鼻海豚的发现率高于中部和北部。问卷调查和随船出海调查表明围网作业发现和误捕小型鲸类的频率要高于拖网 ,同时拖网又高于刺网。调查期间从东山港记录到的误捕致死的 2 0头小型鲸类中 ,6头 (占 30 % )为瓶鼻海豚 ,分别由拖网和围网捕获。需要开展对当地渔民的宣传教育和加强台湾海峡瓶鼻海豚等小型鲸类的种群生物学和保护生物学的研究。