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.     

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.     

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.     

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.     

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.     

ACOUSTIC ECOLOGY OF FORAGING BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS), HABITAT-SPECIFIC USE OF THREE SOUND TYPES

The function(s) of a particular sound can be explored in detail only if the context of its use is well understood. The behavior of the signaler, and the habitat in which that behavior is observed, are two of the most important components of understanding context specific use of a sound. Bottlenose dolphin foraging behavior is often inferred from relatively few behavioral cues that are visible from the surface. To investigate the use of three specific sound types: echolocation, whistles, and pops during foraging, I recorded sound use by animals engaged in a set of previously defined specific foraging behaviors using a system that allowed me to see animals throughout the water column. Lone foraging animals produced all three sounds at significantly higher rates than animals foraging in groups, and the rate of sound production per animal in multi-animal foraging groups did not vary even as the groups reached up to five individuals. Production of echolocation and pops by lone foraging animals accounted for much of the difference. Foraging dolphins also displayed habitat-specific use of particular sound types. They preferentially produced echolocation and pops in the sand habitat and, at least for lone animals, in the seagrass edge habitat.     

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.     

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.     

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.     

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.     

THE OCULAR SECRETIONS OF THE BOTTLENOSE DOLPHIN TURSIOPS TRUNCATUS

A viscous ocular secretion streams from the eyes of bottlenose dolphins, Tursiops truncatuss , removed from the water. The chemical composition and environmentally adaptive implications of this substance continue to perplex curators and scientists. Secretions from three dolphins were examined for deviations from literature values for normal human and terrestrial mammalian tear film components. Osmolality (= 470 mOsm/kg), pH (= 8.44) and glucose (= 26.58 mg/100 ml) were significantly greater than human and terrestrial mammalian tear film values; while lysozyme (= 0.005 mg/ml) and total cholesterol (= 0.17 μg/μ1) were significantly less. Refractive index, sodium ion concentration, and potassium tear to plasma ratios in the dolphin ocular secretion did not deviate significantly from human or terrestrial mammalian tears. Electrophoresis of dolphin tears yielded four major protein bands (probably lactoferrin, serum albumin, teat specific prealbumin, and lysozyme) and high molecular weight glycoproteins similar to human tears. The chemical composition of the T. truncatus ocular secretion suggests that the circumorbital conjunctival gland may be analogous (relative to refractive index, electrolytes, and component protein) to the terrestrial lachrymal gland. At the functional level the T. truncatus ocular secretion may reduce hydrodynamic resistance on the cornea and protect it from bacterial invasion.     

DEVELOPMENT AND SOCIAL FUNCTIONS OF SIGNATURE WHISTLES IN BOTTLENOSE DOLPHINS TURSIOPS TRUNCATUS

ABSTRACT

Bottlenose dolphins Tursiops truncatus produce individually distinctive signature whistles. Dolphins recognize the signature whistles of animals with which they share a social bond. Signature whistles develop within the first few months of life and are stable for a lifetime. Vocal learning appears to play a role in the development of signature whistles in bottlenose dolphins. The signature whistles of most female dolphins and about half of male dolphins differ from those of their mothers. Some dolphin calves born in captivity develop a signature whistle that matches either man-made whistles or those of an unrelated dolphin. Dolphins retain the ability as adults to imitate the whistles of animals with which they share strong individual-specific social relationships, bonds which may change throughout their lifetime. The exceptional imitative abilities of dolphin infants and the retention of this ability in adults may be related to the maintenance of changing individual specific social relationships. Individual recognition by the voice may differ in marine vs terrestrial mammals. Diving marine mammals may not be able to rely upon involuntary voice cues for individual recognition, but rather may require vocal learning to maintain a stable signature as their vocal tract changes shape with increasing pressure during a dive.     

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.     

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.     

SEXUAL DIMORPHISM IN BOTTLENOSE DOLPHINS FROM THE EAST COAST OF FLORIDA

Skulls of 69 bottlenose dolphins (genus Tursiops ) from the Indian/Banana River on the east coast of Florida were examined for evidence of sexual dimorphism. The only sexual dimorphism shown by t -tests on 28 morphological and four meristic skull characters was that males have, on average, more teeth than females in all four arcades. Results of covariance analysis, employed to account for variation in size, indicate minor dimorphism in parietal width of the skull. Twenty body measurements of 29 Tursiops originating in the same area were also analysed for differences between males and females. Statistical results indicate the possibility of sexual dimorphism in the length from the snout to the umbilicus and in flipper width. No evidence was found for differences in overall skull or body length between the sexes.     

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.     

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.     

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.     

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.