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.     

A QUANTITATIVE ASSESSMENT OF DOMINANCE RELATIONS AMONG BOTTLENOSE DOLPHINS

Agonistic behavior of bottlenose dolphins was studied at Brookfield Zoo for nearly 4.5 yr, and dominance relationships were determined using a quantitative technique adapted from primate behavioral research. Dominance relations among dolphins were influenced by the gender of participants. Male dolphins were clearly and consistently dominant to females, and intersexual agonism occurred at moderate rates with seasonal peaks in spring and fall. Dominance relationships among female dolphins were age-ordered and stable, even though agonism among females did occur at uniformly low rates. In contrast, the two males had a changeable dominance relationship in which periods of stability and low-level agonism were interspersed with episodes of intense competition. Zoo-based research revealed patterns of behavior that conformed to current knowledge about bottlenose dolphin social structure. Moreover, research in a zoo setting facilitated development of a quantitative technique that can be used to assess cetacean dominance relationships in field research.     

LOW FREQUENCY NARROW-BAND SOUNDS PRODUCED BY BOTTLENOSE DOLPHINS

We present a new sound type recorded from bottlenose dolphins, Tursiops truncatus , in eastern Australian waters: low-frequency, narrow-band (LFN) harmonic sounds (defined as less than 2 kHz). Most of these sounds were of frequencies less than 1 kHz and were recorded commonly from socializing dolphins. These sounds differ significantly from narrow-band whistles, which are higher in frequency and longer in duration. The absence of these sounds in most studies of the acoustic behavior of bottlenose dolphins may reflect geographic differences in repertoires or result from insufficient sampling. Alternatively, these sounds may have been ignored where the focus of research was on other sound types.     

EXPERIMENTAL RETURN TO THE WILD OF TWO BOTTLENOSE DOLPHINS

In the first scientific experiment of its kind, two young male bottlenose dolphins ( Tursiops truncatus ) were captured in Tampa Bay, Florida, and then returned to the wild at the same locale in October 1990, after two years in captivity. The dolphins' age/sex class and the capture and release site were selected prior to their collection. The ranging and social association patterns of the host community were examined prior to, and, including the two animals, after release. The dolphins remained together for the first month, then began interacting more with other dolphins and less with each other. Within the first year, one dolphin returned to the waters near his capture site and has remained there at least through September 1993. The other dolphin has remained in his original home range at least through June 1996. Observations of each dolphin have shown them to be fully integrated into the local dolphin societies. They displayed typical behavioral, ranging, and social association patterns. Their body condition has been excellent at each observation. They have not been observed interacting with humans. The apparent success of this experiment cannot necessarily be generalized to all potential candidates for return to the wild, but the results can be used to guide future experiments.     

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.     

SURFACE BEHAVIOR OF BOTTLENOSE DOLPHINS IS RELATED TO SPATIAL ARRANGEMENT OF PREY

We tested the hypothesis that spatial arrangement and movements of bottlenose dolphins ( Tursiops truncatus ) are related to the spatial arrangement of their prey. From 65 groups we: (1) classified feeding behavior of dolphins according to spatial arrangement and movements of individuals; (2) assessed spatial arrangement and location of prey from direct observations, numbers of associated seabirds, and echosounder recordings; and (3) related feeding behavior of dolphins to spatial arrangement and location of prey. Four feeding categories were defined from cluster and principal component analyses: (1) moving rapidly with no diving, (2) milling with no diving, (3) diving in several locations, and (4) diving in one location. These feeding categories were related to spatial arrangement and location of prey.     

FLIPPER RUBBING BEHAVIORS IN WILD BOTTLENOSE DOLPHINS (TURSIOPS ADUNCUS)

"Flipper rubbing" behavior was quantitatively analyzed in wild Indo-Pacific bottlenose dolphins ( Tursiops aduncus ) around Mikura Island, Tokyo, Japan. We observed two types of flipper rubbing: (1) F-B rubbing; one dolphin (Rubber) rubbed its flipper over various parts of a partner's (Rubbee) body, and (2) F-F rubbing; both dolphins rubbed each other's anterior flipper edge in alternating shifts. F-B rubbings tended to be initiated by the Rubbee and were terminated by the Rubber. The Rubbee often moved actively its body part that was in contact with the Rubber's flipper, and assumed side-up, upside-down, or other postures while the Rubber remained horizontal in most cases. These facts suggest that the Rubbee engaged in F-B rubbing more actively than the Rubber, and might receive some benefit from the frictional contact during F-B rubbing. Dolphins often switched their roles as Rubber and Rubbee between episodes of flipper rubbing bout. Adults and sub-adults exchanged F-B rubbing and F-F rubbing most often with individuals of the same sex in the same age class. F-B rubbing was frequent in mother-and-calf dyads. Our results suggest that flipper rubbing is an affiliative behavior which could be a quantitative measure of social relationships among individuals of this species in future studies.     

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.     

DIURNAL BEHAVIOR OF FREE-RANGING BOTTLENOSE DOLPHINS IN THE KVARNERIĆ (NORTHERN ADRIATIC SEA)1

The diurnal behavior of a bottlenose dolphin (Tursiops truncatus) community was observed from small inflatable craft between 1987 and 1994. Following a preliminary ad libitum study 11,839 3-min behavioral samples were recorded in 1991-1994. The behavioral budget showed a predominance (about 80%) of activities characterized by long (>30 sec) dives, considered to be largely related to prey search or feeding. Obvious foraging near the surface was observed rarely. The frequent following of trawlers (accounting for 4.6% of the behavioral budget) was indicative of the presence of alternative strategies for finding food. Yearly and seasonal behavioral variation-particularly in feeding-related and travel behaviors-was consistent with the hypothesis of behavioral flexibility as a response to environmental changes and fluctuating prey kind and availability. Yearly shifts in social behavior appeared to be partly influenced by breeding cycles. Groups engaged in feeding-related activities were significantly smaller than traveling or socializing groups, and dramatic interannual group-size shifts seemed to be largely affected by environmental variables, rather than being entirely determined by behavioral activity changes. The remarkable behavioral flexibility of this bottlenose dolphin community may contribute to its survival in the shifting environmental conditions of the northern Adriatic Sea. However, the high proportion of time consistently devoted to feeding-related activities, as compared to other areas, suggests that food resources in the Kvarneri? were not only highly variable but also depleted.     

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.     

THE SHORT-TERM BEHAVIORAL REACTIONS OF BOTTLENOSE DOLPHINS TO INTERACTIONS WITH BOATS IN DOUBTFUL SOUND, NEW ZEALAND

Doubtful Sound is home to one of the southernmost resident populations of bottlenose dolphins ( Tursiops sp.). This population regularly interacts with scenic cruises. During these interactions, dolphins tend to horizontally and vertically avoid vessels, especially when the behavior of these vessels is intrusive. This study aimed at understanding the behavioral reactions of individuals to these interactions that lead to the disruption of the school's behavioral state. Observing the behavioral events performed by individuals during an interaction can help define the short-term reactions elicited by the boat presence. I recorded the behavioral events performed by all individuals of focal schools. The frequency of occurrence of all events was compared depending on the presence of vessels, their behavior, and the behavioral state of the focal school. Dolphins tended to perform more side flops while interacting with powerboats, a behavior which may be used as a non-vocal communication tool. Moreover, the movement of dolphins became more erratic during interactions with all types of vessels. These effects increased when the boats were more intrusive while interacting. This study shows that the impact of interaction with boats can be minimized if the vessels respect the guidelines in place.     

SOCIAL ECOLOGY OF BOTTLENOSE DOLPHINS IN THE KVARNERIĆ (NORTHERN ADRIATIC SEA)

A bottlenose dolphin community was studied from small inflatable craft from 1987 to 1994 in a relatively large area (about 800 km²) east of the islands of Loˇsinj and Cres, northern Adriatic Sea. A total of 106 individuals were photoidentified based on natural permanent marks on their dorsal fins. Most of the dolphins were resighted on a regular basis, indicating a high level of year-round site fidelity, although their range was evidently greater than the chosen study area. Dolphin density was highly variable and considerably lower than for most well-known bottlenose dolphin communities. Groups averaged seven individuals, with a mode of two. Groups entirely composed of adults were the smallest, groups with calves the largest. Group fluidity was high, seasonal and yearly changes in mean group size being also considerable. Summer was the peak calving season, with a striking variation in the number of births on alternate years. Poor evidence of shark predation was found. The social organization of this dolphin community seemed to be highly flexible, possibly as an adaptation to cope with environmental changes as well as with a limited and variable availability of prey.     

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.     

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.     

GEOGRAPHIC VARIATION IN RATES OF VOCAL PRODUCTION OF FREE-RANGING BOTTLENOSE DOLPHINS

Echolocation and whistle production, group sizes, and activities of free-ranging bottlenose dolphins were compared across four regions (Wilmington, NC Intracoastal Waterway [ICW]; Wilmington coastline; Southport, NC coastline; and Sarasota, FL inshore waters). Number of whistles and echolo-cation bouts differed significantly across sites. Dolphins whistled significantly more in Southport than in the other sites, independent of group size. Unlike at the other sites, dolphin vocalizations in Southport did not vary significantly across activities; this difference may be due to the fact that Southport animals were often found behind shrimp-trawling vessels, which may affect their behavior. Resident Sarasota dolphins vocalized significantly less than dolphins at the NC sites. At most sites, echolocation production per dolphin decreased as group size increased, supporting the idea that echolocation information is shared. In the ICW and Sarasota, echolocation production per dolphin was highest while feeding, indicating that echolocation is used in foraging. At all sites but Southport, whistle production per dolphin was highest while socializing, indicating that whistles are used in communication. Overall, these data show that dolphins have different vocal and activity patterns at different sites; thus, caution should be used when extrapolating results from one study site to another.     

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.     

RAY ENCOUNTERS AS A MORTALITY FACTOR IN ATLANTIC BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS)

Ray spines were found associated with various tissues on necropsy of seven Atlantic bottlenose dolphins ( Tursiops truncatus ). In at least six cases they were considered a major factor in the death of the animal. Three mature Tursiops had ray spines located in the lung parenchyma. In two other dolphins the ray spine punctured the lateral chest wall without entering the lung tissue. A sixth Tursiops contained a ray spine located in the liver tissue, while a seventh animal had a ray spine lodged in the pancreas. Five of the dolphins were wild individuals and two had been in captivity for more than one year. These findings indicate that beached dolphins and dolphins in captivity with a chronic history of illness should be examined for the presence of ray spines. Caution should be used in maintaining mixed exhibits of dolphins with stingray species.     

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.