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.     

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.     

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.     

CHARACTERIZATION OF A BOTTLENOSE DOLPHIN (TURSIOPS TRUNCATUS) KIDNEY EPITHELIAL CELL LINE

Abstract: An epithelial cell line, Carvan dolphin kidney (CDK), isolated from a prematurely born female bottlenose dolphin, Tursiops truncatus , exhibited growth characteristics not previously reported for cetacean cells in culture. CDK cells were cytokeratin positive and demonstrated a maximum doubling time of 1.31 days, with plating and colony forming efficiencies approaching 100% for the early population doublings. Despite an unusually efficient colony-forming ability and rapid growth, these cells were neither transformed nor immortal, displaying normal contact inhibition, anchorage dependence, and the requirement for high concentrations of fetal bovine serum in the growth medium. CDK cells exhibited age-dependent changes in growth rate, colony-forming efficiency, and cytoplasmic profile, and showed a finite lifespan of about 50 population doublings and a stable 2N = 44 karyotype which correlates with previously reported cytogenetic analyses. Velocity sedimentation analysis showed that CDK cells contained nuclear aryl hydrocarbon receptor (Ah), indicating their potential to be induced for cytochrome P450. These data suggest that CDK cells may have utility as an in vitro toxicological model for evaluating hydrocarbon contaminant effects on Tursiops truncatus , a protected marine mammal.     

INTERACTIONS BETWEEN THE INDIAN RIVER LAGOON BLUE CRAB FISHERY AND THE BOTTLENOSE DOLPHIN, TURSIOPS TRUNCATUS

Anecdotal reports from blue crab fishermen in the Indian River Lagoon (IRL), Florida suggested that bottlenose dolphins ( Tursiops truncatus ) followed their boats, and stole bait fish from crab pots soon after they were deployed. To investigate these reports, we made biweekly observations from IRL commercial crab boats from January 1998 to January 1999 (670 h). Only 2.8% of the 18,891 crab pots surveyed revealed evidence of dolphin/crab pot interaction. Dolphin interactions included: (1) begging at boats, (2) feeding on discarded bait fish, (3) engaging in crab pot tipping behavior, and (4) dolphin mortality from crab pot float line entanglement. Overall, 16.6% of the 1,296 dolphins sighted interacted with fishing boats. Seasonal trends were evident, with fishery interactions peaking in the summer. Crab pot interactions ranged from 0% to 36% of the traps checked daily. Different methods of securing the bait-well door and the role of trap locality were tested using a replicated experimental design. Results showed significant differences in successful bait removal by dolphins, among the degrees of door security ( P < 0.001) and between trap location ( P < 0.01). Thus, increased door security may help to reduce the negative impacts to the fishery and dolphins involved.     

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.     

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.     

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.     

NO EVIDENCE OF ACCOMMODATION IN THE EYES OF THE BOTTLENOSE DOLPHIN, TURSIOPS TRUNCATUS

Bottlenose dolphins ( Tursiops truncatus ) are aquatic mammals that must come to the surface to breathe. As a result, it might be expected that their eyes are adapted for both aerial and underwater vision. Earlier studies suggest that dolphins are emmetropic ( i. e. , focused at infinity) in water, and in some cases, emmetropic in air, although the mechanisms that permit these animals to see well in both media are not well understood. Nor is it known whether they can accommodate to focus sharply on objects at different distances. We employed video photoretinoscopy to investigate the possibility of an active accommodative mechanism in the eyes of the bottlenose dolphin in water. Measurements of the refractive state in water indicated near emmetropia for two individuals and slight myopia (nearsightedness) for the third individual. No clear cases of accommodation were observed underwater in any of the subjects examined. Vision underwater may be used to supplement echolocation. If so, such a role might not require an accommodative mechanism.     

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.     

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.     

TEMPORARY THRESHOLD SHIFTS AFTER NOISE EXPOSURE IN THE BOTTLENOSE DOLPHIN (TURSIOPS TRUNCATUS) MEASURED USING EVOKED AUDITORY POTENTIALS

The time course of recovery from temporary threshold shift (TTS) was measured in a bottlenose dolphin, Tursiops truncatus , using an evoked-potential procedure. The envelope-following response (EFR), which is a rhythmic train of auditory brainstem responses (ABR) to sinusoidally amplitude-modulated tones, was used as an indicator of the sound reception by the animal. Variation of the intensity of the stimulus allowed us to measure the animal's hearing via EFR thresholds. During each session, following an initial measure of threshold, the trained animal voluntary positioned itself within a hoop 1 m underwater while a 160 dB re 1 μPa noise of a 4–11 kHz bandwidth was presented for 30 min. After the noise exposure, thresholds were measured again at delays of 5, 10, 15, 25, 45, and 105 min. Measurements were made at test frequencies of 8, 11.2, 16, 22.5, and 32 kHz. The maximum TTS occurred 5 min after exposure and rapidly recovered with a rate of around 1.5 dB per doubling of time. TTS occurred at test frequencies from 8 to 16 kHz, with the maximum at 16 kHz. TTS was negligible at 22.5 kHz and absent at 32 kHz.     

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.     

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.     

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.     

AGE STRUCTURE AND GROWTH OF THE BOTTLENOSE DOLPHIN (TURSIOPS TRUNCATUS) FROM STRANDINGS IN THE MISSISSIPPI SOUND REGION OF THE NORTH-CENTRAL GULF OF MEXICO FROM 1986 TO 2003

Despite their high abundance and wide distribution, little is known about the historical or current growth and age structure of coastal bottlenose dolphins ( Tursiops truncatus ) in the north-central Gulf of Mexico. Between 1986 and 2003, teeth were collected from bottlenose dolphins stranded on the mainland coast of Mississippi and the adjacent barrier islands. Bottlenose dolphin strandings occurred year round, peaking in March and April. Neonate strandings also peaked during these 2 mo. Age estimates were obtained from 111 animals by reading the growth layer groups in the dentine layer of the teeth. The ages ranged from <1 yr to 30 yr of age. The two-stage Laird–Gompertz growth model was fitted to the total length and age data. On the basis of this model, the asymptotic lengths were estimated at 250 cm for females and 255 cm for males. The length at birth estimates were 98–103 cm for females and 100–107 cm for males. These lengths are similar to those of bottlenose dolphin populations from other Gulf of Mexico areas and from the North Atlantic Ocean along the southeastern United States.     

HABITAT USE PATTERNS AND RANGES OF THE BOTTLENOSE DOLPHIN IN THE GULF OF CALIFORNIA, MEXICO

Abstract: I studied behavior and range patterns of individual bottlenose dolphins during 1984 in the mid-eastern Gulf of California, Mexico. Dolphin sighting rate was significantly higher in areas close to estuary mouths, 0.306 sightings per hour compared with 0.155 sightings per hour in areas distant from estuary mouths. Dolphins used these estuarine areas to feed; 61% of all behavior observed near estuaries was feeding as compared with 23% elsewhere. Traveling comprised 61% of all behavior observed in areas distant from estuary mouths. Estuaries are sites of large concentrations of nutrients which support great numbers of filter-feeding zooplankton and fish. Bottlenose dolphins may specialize on esmarine prey, or they may feed in estuarine areas simply because of the abundance of potential prey that these systems support. In either case, data on relative numbers, distribution patterns, behavior and diet indicate that this is a general trend in habitat use for many coastal populations of this species in the Pacific and Atlantic.
Ranges of a few individuals spanned a minimum of 65 km of coastline, and animals were not permanent residents of a monitored bay. In contrast, dolphins off the coast of Sarasota, Florida, have been reported to be year-round residents with smaller ranges. This difference in degree of site fidelity may be related to habitat differences. The west coast of Florida is dotted with numerous and large estuarine systems which may host permanent prey populations and support resident groups of dolphins. The Gulf of California coastline contains few estuaries; most are small and perhaps support prey resources which are ephemeral, requiring dolphins to range over larger distances in search of food.     

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.     

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.     

CROSS-REACTIVITY OF ANTIBODIES TO HUMAN ANTIGENS WITH TISSUES OF THE BOTTLENOSE DOLPHIN, TURSIOPS TRUNCATUS, USING IMMUNOPEROXIDASE TECHNIQUES

Cross-species reactivity of 53 antibodies developed against human antigens was evaluated in the bottlenose dolphin Tursiops truncatus , using immunoperoxidase techniques, Strong cross-reactivity was demonstrated with antibodies against intermediate filaments, most hormones, but few leukocyte antigens. S100, NSE, F8RA, Factor XIIIa, and lactalbumin antibodies reacted, while HMB45, EMA, PLAP, and PSA did not. Polyclonal antibodies are much more likely to cross-react than are monoclonal antibodies. Twenty-four of 30 (80%) of polyclonals cross-reacted, while only 10 of 23 (43.5%) monoclonals cross-reacted. Some antigens are demonstrated only after using a special technique to unmask the antigen.