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.     

POPULATION GENETIC STRUCTURE OF COASTAL BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) IN THE NORTHERN BAHAMAS

Population substructure has important implications for a species' ecology and evolution. As such, knowledge of this structuring is critical for the conservation and management of natural populations. Among marine mammals, many examples exist of species that enjoy a broad geographical distribution, yet are characterized by fine‐scale population subdivisions. Coastal bottlenose dolphins have been studied extensively in a few regions globally, and these studies have highlighted a great diversity in both social strategies and demographic isolation. Here we use molecular genetic markers to examine the degree of population subdivision among three study sites separated by less than 250 km on Little Bahama Bank in the northern Bahamas. Mitochondrial DNA (mtDNA) sequence variation and microsatellite genotypes were used to assess partitioning of genetic variance among 56 individually recognized coastal ecotype bottlenose dolphins. Although resolved levels of genetic differentiation suggest gene flow among the three study sites, both nuclear and mitochondrial data indicate a significant degree of subdivision within the Little Bahama Bank population, and sex‐based analyses suggest that patterns of dispersal may not be strictly biased toward males. These results corroborate the site fidelity documented through long‐term photo‐identification studies in the NE Bahamas, and highlight the need to consider independent subpopulation units for the conservation and management of coastal bottlenose dolphins in the Bahamas.     

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.     

COASTAL BOTTLENOSE DOLPHINS FROM SOUTHEASTERN AUSTRALIA ARE TURSIOPS ADUNCUS ACCORDING TO SEQUENCES OF THE MITOCHONDRIAL DNA CONTROL REGION

Sequence analysis of the mitochondrial DNA control region was used to clarify the taxonomic status of two coastal bottlenose dolphin populations from southeastern Australia currently classified as Tursiops truncatus . A 368-bp segment of the control region of 57 biopsy-sampled, photo-identified dolphins of Jervis Bay and Port Stephens was compared to published sequences of T. truncatus and T. aduncus from different oceanic regions. Sequence divergence between haplotypes from southeastern Australia and T. aduncus was much lower than that from T. truncatus . Analyses using two different methods of phylogenetic reconstruction unambiguously placed all haplotypes from southeastern Australia in a group composed exclusively of T. aduncus . The results strongly indicated that these two bottlenose dolphin populations belong to T. aduncus , extending the range of the species to subtropical waters of the Western South Pacific Ocean.     

MOTHER-INFANT SPATIAL RELATIONS IN CAPTIVE BOTTLENOSE DOLPHINS,TURSIOPS TRUNCATUS

The prolonged nursing period and strong, extended mother-infant bond observed among bottlenose dolphins may reflect social and physical ontogeny critical for infant survival. This study was conducted to quantify ontogentic changes in mother-infant contact time and the amount of time infants spent in specific spatial states with their mothers from birth to age 12 mo. These behaviors were studied through a systematic, longitudinal study of six mother-infant pairs of captive bottlenose dolphins from three different social groups. There was a significant decrease in the time infants spent with their mothers (logistic regression, P < 0.001), following the general mammalian pattern of increasing independence with age. When with their mothers, the probability that infants would be found in “echelon” position, flanking the mother, decreased as the calf aged (logistic regression, P <0.001), possibly due to anatomical and hydrodynamic factors. The probability that infants would be found in “infant” position, underneath the mother, increased with calf age (logistic regression, P < 0.001). Results obtained in this study are consistent with similar studies of wild bottlenose dolphin mother-infant pairs, indicating a suite of ontogenetically comparable behaviors between wild and captive bottlenose dolphins.     

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.     

POPULATION STRUCTURE OF THE BOTTLENOSE DOLPHIN (TURSIOPS TRUNCATUS) AS DETERMINED BY RESTRICTION ENDONUCLEASE ANALYSIS OF MITOCHONDRIAL DNA

Abstract: Restriction fragment length polymorphisms of mitochondrial DNA (mtDNA) were used to test for population subdivision in the bottlenose dolphin (Tursiops truncatus). Atlantic and Pacific dolphin mtDNA samples exhibited distinctly different haplotypes (approximately 2.4% sequence divergence), indicating a lack of gene exchange. Within the Atlantic Ocean, mtDNA samples from the Gulf of Mexico and the Atlantic Coast were also found to be distinct, with a sequence divergence of approximately 0.6%. The Atlantic Coast–Gulf of Mexico dichotomy is consistent with patterns of genetic variation from other marine and coastal organisms from this region, and supports the hypothesized role of bio-geographic events in promoting the divergence of these and other forms. Regional differentiation was identified along the Atlantic Coast, whereas low sequence divergences among haplotypes and consistent haplotype frequencies across populations suggested considerable gene exchange among Gulf of Mexico populations. A highly divergent haplotype found in two individuals from two localities in the Gulf of Mexico is best explained by dispersal from either a distinct offshore Gulf stock or an unsampled Atlantic Coast stock. Additional samples are required to test for the existence of a distinct offshore race and, if it exists, to identify its distribution and contribution to population structure.     

HIGH PERFORMANCE TURNING CAPABILITIES DURING FORAGING BY BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS)

Large predators should have difficulty catching small prey because small animals demonstrate greater maneuverability and agility compared to large animals. The ability of a predator to capture small prey indicates locomotor strategies to compensate for inequities in maneuverability. Bottlenose dolphins ( Tursiops truncatus ) in Sarasota Bay, Florida feed on fish at least one order of magnitude smaller than themselves. To examine the locomotor strategies involved in prey capture, the foraging movements of these dolphins were videotaped from overhead using a remotely-controlled camera suspended from a helium-filled aerostat, which was tethered to an observation vessel. Dolphins were observed to rapidly maneuver during chases of fish in open water or around patches of rooted vegetation. Video analysis of the chase sequences indicated that the dolphins could move the rostrum through small radius turns with a mean value of 0.20 body lengths and with a minimum value of 0.08 body lengths. Mean rate of turn was 561.6°/sec with a maximum rate measured at 1,372.0°/sec. High turning rates with small turning radii were primarily the result of maneuvers in which the dolphin rolled 90° and rapidly flexed its body ventrally. The ability of dolphins to change body orientation in multiple rotational axes provides a mechanism to reduce turning radius and increase turning rate to catch small, elusive prey.     

DEMOGRAPHIC PARAMETERS OF INDO-PACIFIC BOTTLENOSE DOLPHINS (TURSIOPS ADUNCUS) AROUND MIKURA ISLAND, JAPAN

Long-term identification surveys were conducted using video records on Indo-Pacific bottlenose dolphin ( Tursiops aduncus ) around Mikura Island, Tokyo, Japan. On 246 surveys, 3,996 min of video were recorded during each summer season from 1994 to 2001. Two hundred and twenty individuals were observed, including juveniles and neonates, that were not cataloged, but could be identified by association with their mothers. Over the eight-year study period, the total number of identified and cataloged individuals was 169. The annual number of newly identified dolphins stabilized at ∼ 10 after 1995. In the middle of each research season, a plateau in the number of reidentified individuals was reached. Most of the dolphins were observed frequently–percentages of reidentified dolphins in adjoining years were greater than 86% and 62 individuals were seen consecutively during the eight years. There were no significant differences in the sex ratio between years. The sex ratio was skewed significantly towards male for subadults and towards female for adults. The survival rate of one-year-old calves was 86.7%, and the mean age at weaning was 3.5 yr. The calving interval averaged 3.4 yr among adult females. The mean annual birth, fecundity, and recruitment rates were 0.071, 0.239, and 0.068, respectively. Mikura Island is located in the pelagic ocean with no protected bay area existing around the island. Dolphins do not regularly inhabit any other locations besides Mikura within the Izu island chain. Thus, this island may be important core habitat of these dolphins.     

EFFECTS OF WATERCRAFT NOISE ON THE ACOUSTIC BEHAVIOR OF BOTTLENOSE DOLPHINS, TURSIOPS TRUNCATUS, IN SARASOTA BAY, FLORIDA

Watercraft may provide the greatest source of anthropogenic noise for bottlenose dolphins living in coastal waters. A resident community of about 140 individuals near Sarasota, Florida, are exposed to a vessel passing within 100 m approximately every six minutes during daylight hours. I investigated the circumstances under which watercraft traffic may impact the acoustic behavior of this community, specifically looking for short-term changes in whistle frequency range, duration, and rate of production. To analyze whistles and received watercraft noise levels, acoustic recordings were made using two hydrophones towed from an observation vessel during focal animal follows of 14 individual dolphins. The duration and frequency range of signature whistles did not change significantly relative to vessel approaches. However, dolphins whistled significantly more often at the onset of approaches compared to during and after vessel approaches. Whistle rate was also significantly greater at the onset of a vessel approach than when no vessels were present. Increased whistle repetition as watercraft approach may simply reflect heightened arousal, an increased motivation for animals to come closer together, with whistles functioning to promote reunions. It may also be an effective way to compensate for signal masking, maintaining communication in a noisy environment.     

LOCAL ABUNDANCE AND DISTRIBUTION OF BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) IN THE NEARSHORE WATERS OF VIRGINIA BEACH,VIRGINIA

We investigated patterns of abundance and distribution for coastal migratory Atlantic bottlenose dolphins (Tursiops truncatus) that appear seasonally in the nearshore waters of Virginia Beach, Virginia. The study was conducted along 24 km of shoreline at the southern point of the Chesapeake Bay mouth from April 1994 to March 1995. This is the first study to investigate the relationship between the abundance of coastal migratory dolphins and factors that might affect their movement. A profile analysis of variance revealed significant differences in local abundance and distribution throughout the year. Dolphin number was positively correlated with water temperature and not correlated with photoperiod. Although prey distribution and abundance are two factors thought to affect dolphin presence, in this study the relationship between these two factors and dolphin abundance was unclear. Greater numbers of dolphins were found in the ocean section of the study area. However, significantly higher ratios of neonatal dolphins were observed in the bay section, suggesting the bay serves as a nursery area. The observed relationship between local dolphin abundance and environmental factors in Virginia may provide insight into dolphin distribution and migration along the Atlantic coast of the United States.     

中国水域瓶鼻海豚的mtDNA控制区序列变异性分析

测定了30头中国水域瓶鼻海豚（Tursiops sp.)mtDNA控制区5′端424bp的序列,结合已发表的中国水域其它瓶鼻海豚的mtDNA控制区序列,共发现54个变异位点,定义了37种单元型。中国水域瓶鼻海豚的两个形态型之间没有共享单元型,且具有8个鉴别位点。基于最大似然法和邻接法的系统发生分析均把单元型聚类为分别代表两个形态型的支系。形态型之间的核苷酸歧异度为5.58%,超过了其它海豚类种间的序列歧异水平,支持把这两个形态型划分为两个独立的种,即T.truncatus和T.aduncus的观点。虽然两种瓶鼻海豚的分布区在台湾海峡一带出现重叠,但相互之间缺乏基因流动,提示两者可能已出现了显著的生殖隔离。     

INCREASED AVOIDANCE OF SWIMMERS BY WILD BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) DUE TO LONG-TERM EXPOSURE TO SWIM-WITH-DOLPHIN TOURISM

The responses of wild, non-provisioned bottlenose dolphins ( Tursiops truncatus ) to swim attempts from commercial swim-with-dolphin tour boats were systematically observed during two research periods: 1994–1995 and 1997–1998. A total of 255 groups of dolphins was encountered during boat-based surveys and 36% ( n = 93) were exposed to at least one swim attempt. The operators' success with swim attempts, defined as at least one dolphin milling within 5 m of at least one swimmer, decreased from 48% in 1994–1995 to 34% in 1997–1998, and avoidance responses to swimmers increased from 22% to 31%. Dolphin response was found to vary according to swimmer placement. The greatest increase in avoidance occurred when swimmers were placed in the dolphins' path of travel. Based on sighting records of 266 individually identified dolphins, it was estimated that an average dolphin was exposed to 31 swim attempts per year. This level of exposure suggests that individual dolphins have, with cumulative experience, become sensitized to swim attempts. When a swim attempt was successful, on average it involved 19% of the group. Age-class differences in interaction rates showed that juveniles were significantly more likely to interact with swimmers than adults. This study highlights the importance of longitudinal studies in evaluating human impact and suggests the urgent need for similar studies of potential human impact on other toothed cetaceans.     

A RESIDENT POPULATION OF INDO-PACIFIC BOTTLENOSE DOLPHINS (TURSIOPS ADUNCUS) IN AMAKUSA, WESTERN KYUSHU, JAPAN

Residency patterns and population size were examined for Indo-Pacific bottlenose dolphins ( Tursiops aduncus ) in the coastal waters of Amakusa-Shimoshima Island, western Kyushu, Japan, from summer 1994 to fall 1998. A total of 203 individuals were identified. The number of newly identified individuals barely increased after the first 1.5 yr. The identified individuals in one season were frequently resighted with percentages of mostly over 60% during the subsequent seasons. Most of the dolphins off Amakusa were yearround residents, although the total range of their habitat was unknown. The population size estimated by the mark-recapture technique was 218 individuals with a CV of 5.41% in 1995–1997. The group commonly consisted of more than 100 individuals. The large group size was thought to be a response to feeding on schooling fishes.     

SATELLITE-MONITORED MOVEMENTS AND DIVE BEHAVIOR OF A BOTTLENOSE DOLPHIN (TURSIOPS TRUNCATUS) IN TAMPA BAY, FLORIDA

An adult, female bottlenose dolphin ( Tursiops trucncatus ) was radio tagged and monitored via satellite-based Argos receivers for 25 d from 28 June to 23 July 1990, in Tampa Bay, Florida. A total of 794 transmissions were obtained during 106 satellite passes. A mean of 3.9 (SE = 0.24) locations/day were determined by Service Argos and showed the animal remained in the bay, usually close to the southeastern shore. The dolphin moved at least 581 km at a minimum mean speed of 1.2 (SE = 0.1) km/h. Data from 63, 922 dives were recorded. The animal spent an average of 87.1 (SE = 0.6)% of the time submerged, with a mean dive duration of 25.8 (SE = 0.5) sec. Mean dive duration differed significantly between four periods of the day, as did the mean percent of time spent submerged. During the early morning the animal spent more time at the surface, averaged shorter dives, and was submerged less than other times of day. This is the first study to demonstrate die1 dive cycles in a bottlenose dolphin. Four months after tag loss, the dolphin was photographed with no evidence of necrosis or disfigurement of the dorsal fin. Satellite telemetry was demonstrated as an effective means of documenting the movements and dive behavior of a small inshore cetacean.     

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.     

GROWTH OF BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) FROM THE INDIAN RIVER LAGOON SYSTEM, FLORIDA, U. S. A.

Gompertz growth models were fitted to total lengths and ages from tooth sections of 199 stranded bottlenose dolphins ( Tursiops truncatus ) from the Indian River Lagoon system, eastern Florida. Based on the model, dolphins from this population are estimated to be born at 119 cm and reach asymptotic length at 250 cm. No apparent pubescent growth acceleration was noted for either sex. Males appeared to grow to slightly longer lengths than females. There were small size differences between Indian River dolphins and those in Texas and Sarasota, indicating general size similarities between North Atlantic and Gulf of Mexico bottlenose dolphins stocks.     

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.     

A MODEL LIFE TABLE FOR BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) FROM THE INDIAN RIVER LAGOON SYSTEM, FLORIDA, U.S.A.

Data gathered from 220 stranded bottlenose dolphins ( Tursiops truncatus ) in the Indian River Lagoon system, Florida, were used to derive a life table. Survivorship curves were fit to the data using Siler's competing-risk model and a maximum likelihood approach. Population growth was estimated to be between r = 0.0 and 0.046 based on the observed numbers of stranded dolphins. Variance in survival rates was estimated using an individual-based, age-structured population projection model. We estimate that the overall annual mortality rate for this population was 9.8% per year. Sex-specific differences in survivorship were apparent with females outliving males. The overall mortality curve resembles that of other large mammals, with high calf mortality and an exponentially increasing risk of senescent mortality. The inclusion of live-capture removals of individuals from this population did not significantly affect the estimation of survival parameters for most age classes.     

A BIOPSY SYSTEM FOR SMALL CETACEANS: DARTING SUCCESS AND WOUND HEALING IN TURSIOPS SPP.

Together with PAXARMS (NZ), we developed a biopsy system for small cetaceans and tested it on four populations of bottlenose dolphins ( Tursiops spp.). The system consists of a modified 0.22 caliber rifle, and biopsy darts made out of polycarbonate with stainless steel biopsy tips. Animals were darted at a range of 2–15 m while travelling parallel to the vessel. Overall sampling success for obtaining biopsy samples when an animal was struck ranged from 96.6% to 100% in the four populations. However, hit rate varied for the four different populations. We did not observe a significant difference in strength of the reaction to the darting procedure when an animal was hit or missed, both among and within populations. Data from one population showed no significant difference in the reaction to biopsy sampling by four different age-sex classes. The only factor that had a significant influence on darting success was the hit location. Furthermore, we observed a significant positive correlation between the size of the sample obtained and the reaction to biopsy sampling. Biopsy samples were sufficient for microsatellite and d-loop analysis in 95.8% and for genetic sexing in 99% of all cases. In animals that we observed on a daily basis, wounds were healed after approximately 23 d.