Population structure of island-associated dolphins: Evidence from photo-identification of common bottlenose dolphins (Tursiops truncatus) in the main Hawaiian Islands

Management agencies often use geopolitical boundaries as proxies for biological boundaries. In Hawaiian waters a single stock is recognized of common bottlenose dolphins, Tursiops truncatus , a species that is found both in open water and near-shore among the main Hawaiian Islands. To assess population structure, we photo-identified 336 distinctive individuals from the main Hawaiian Islands, from 2000 to 2006. Their generally shallow-water distribution, and numerous within-year and between-year resightings within island areas suggest that individuals are resident to the islands, rather than part of an offshore population moving through the area. Comparisons of identifications obtained from Kaua'i/Ni'ihau, O'ahu, the "4-island area," and the island of Hawai'i showed no evidence of movements among these island groups, although movements from Kaua'i to Ni'ihau and among the "4-islands" were documented. A Bayesian analysis examining the probability of missing movements among island groups, given our sample sizes for different areas, indicates that interisland movement rates are less than 1% per year with 95% probability. Our results suggest the existence of multiple demographically independent populations of island-associated common bottlenose dolphins around the main Hawaiian islands.     

Progress report: Results of a long-range captive breeding program for the bottlenose dolphin,Tursiops truncatus and Tursiops truncatus gilli

There have been 36 bottlenose dolphins born in breeding colonies at Sea World, California, and Sea World, Florida, from 1978–1985. Significant features of this successful reproductive program are construction of a compatible breeding colony, early hormonal detection of pregnancy, pre- and postnatal association of inexperienced mothers with experienced females and their calves, and minimized handling of the mothers and calves until the calves are over 1 year of age. Inclusion in the breeding colonies of males old enough to be effective breeders is stressed. Females in the colonies have successfully bred from 8–9 years of age to 23–24 years of age. Calving intervals in the colonies vary from 2 to over 3 years. Calves are born all year round, with some peaking in calving activity in the spring and fall months. Serum progesterone levels greater than 6,000 pg/ml, maintained over a 4–6-week period, are considered indicative of pregnancy. Progesterone levels vary from less than 10,000 to over 50,000 pg/ml during gestation, averaging 25,000 pg/ml. The need for further study of variation in the pattern of progesterone levels during pregnancy is emphasized. A gestation period for Tursiops of 11.5 to 12 months is consistent with our cumulative progesterone data.     

Estimating the impact of interactions between bottlenose dolphins and artisanal fisheries around the Balearic Islands

Interactions between marine mammals and fisheries are a growing problem, and effective management requires assessment of the factors driving the interaction and of the impacts on fisheries. We used data from interactions between artisanal fisheries and bottlenose dolphins around the Balearic Islands to assess these factors and impacts. Observers collected data during 1,040 fishing operations over 3 yr. Location and year were important factors affecting interaction probability, with some areas showing large increases over the study period. We estimated the combined cost of catch loss and net damage as 6.5% of the total catch value (95% CI ?12.3%, ?1.6%), and the annual loss to be 3.4% (95% CI ?6.5%, ?0.1%) of the total catch by weight. This weight equates to the dietary needs of ～12 dolphins (95% CI 0.2, 22), suggesting the fishery is not a vital food source for the dolphin population. Two dolphins died through entanglement during the observed fishing operations. We observed 3% of the total fishing activity, by weight, in 2003; scaling up this mortality directly suggests that as many as sixty dolphins may be dying in nets each year. This interaction likely has serious conservation implications for the dolphin population.     

Differences in acoustic signals from Delphinids in the western North Atlantic and northern Gulf of Mexico

Whistle characteristics were quantitatively compared between both geographically separated and neighboring populations of Atlantic spotted dolphins (Stenella frontalis), bottlenose dolphins (Tursiops truncatus), and pilot whales (Globicephala spp.) in U.S. waters to evaluate if intraspecific acoustic differences exist between groups. We compared nine whistle characteristics between continental shelf and offshore Atlantic spotted dolphins in the western North Atlantic and between northern Gulf of Mexico and western North Atlantic bottlenose dolphins and pilot whales using discriminant analysis. Offshore Atlantic spotted dolphin whistles were significantly different (Hotelling's T², P= 0.0003) from continental shelf whistles in high frequency, bandwidth, duration, number of steps, and number of inflection points. Atlantic bottlenose dolphin whistles were significantly different (Hotelling's T², P < 0.0001) from those in the Gulf of Mexico in duration, number of steps, and number of inflection points. There was no significant difference between pilot whale whistles in the two basins. The whistle differences indicate acoustic divergence between groups in different areas that may arise from geographic isolation or habitat separation between neighboring but genetically distinct populations of dolphins. This study supports the premise that acoustic differences can be a tool to evaluate the ecological separation between marine mammal groups in field studies.     

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

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

     

Epidemiology of tattoo skin disease in captive common bottlenose dolphins (Tursiops truncatus): Are males more vulnerable than females?

Clinical and epidemiological features of tattoo skin disease (TSD) are reported for 257 common bottlenose dolphins held in 31 facilities in the Northern Hemisphere. Photographs and biological data of 146 females and 111 males were analyzed. Dolphins were classified into three age classes: 0–3 years, 4–8 years, and older than 9 years. From 2012 to 2014, 20.6% of the 257 dolphins showed clinical TSD. The youngest dolphins with tattoo lesions were 14 and 15 months old. TSD persisted from 4 to 65 months in 30 dolphins. Prevalence varied between facilities from 5.6% to 60%, possibly reflecting variation in environmental factors. Unlike in free-ranging Delphinidae, TSD prevalence was significantly higher in males (31.5%) than in females (12.3%). Infection was age-dependent only in females. Prevalence of very large tattoos was also higher in males (28.6%) than in females (11.1%). These data suggest that male T. truncatus are more vulnerable to TSD than females, possibly because of differences in immune response and susceptibility to captivity-related stress.     

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.     

Advancing the use of morphometric data for estimating and managing common bottlenose dolphin (Tursiops truncatus) mass

Knowledge of a dolphin's body mass is central to establishing body condition, comparing across individuals, and designing successful management programs. In the present study, sex‐specific prediction equations for estimating body mass were generated from morphometrics (i.e., length and girth) and ages of bottlenose dolphins residing under professionally managed care. Measurements of wild dolphins in Sarasota Bay, Florida, were used to generate sex‐specific body mass reference ranges. Gompertz growth models were fitted to length measurements and age to compare growth across populations. From the regression analyses, the body mass of managed females (R² = 0.937), managed males (R² = 0.953), wild females (R² = 0.979), and wild males (R² = 0.972) were predicted with high levels of accuracy. Managed adults had similar or longer asymptotic lengths compared to their wild conspecifics. To apply this information, ZooMorphTrak, a mobile software application, was developed to provide a new resource for management. The “Approximate” feature was designed to approximate body mass based on user inputs of individual morphometrics. The “Management” feature compared a managed dolphin's known body mass with respect to body mass reference ranges generated from wild dolphins. ZooMorphTrak, developed by the Chicago Zoological Society, is available for download at http://itunes.apple.com .     

Modeling Response of Target and Nontarget Dolphins to Biopsy Darting

Abstract: Remote biopsy sampling is a valuable technique to collect data on free-ranging cetaceans. Numerous researchers have evaluated effects of biopsy darting on the target animal, but biopsy sampling may also have unintentional effects on nearby nontarget animals. We evaluated reactions of target and nontarget animals during remote biopsy sampling conducted on bottlenose dolphins (Tursiops truncatus) in August 2004 off Folly Beach, South Carolina, and Jekyll Island, Georgia, USA, and in February-March and July-August 2005 off Hilton Head, South Carolina. We examined external factors that might influence response of the target animal as well as other individuals in the group. Target animals reacted to biopsy attempts 90.9% (n = 475) of the time. Of these, 97.5% (n = 432) of reactions were to biopsy hits, and 84.4% (n = 320) of all hits had reactions recorded as low. Nontarget animals reacted 21.9% (n = 1,158) of the time; interanimal distance from target animal, season, sea state, and depth affected reactions of nontarget animals. Overall, however, biopsy effects appear to be minimal. We present a statistical tool for identifying factors that elicit greater reactions. This model could be adapted for species or locations, as needed, to identify factors that could influence effects of biopsy sampling.     

Ontogenetic diet changes in bottlenose dolphins (Tursiops truncatus) reflected through stable isotopes

The ability of stable isotope analysis to provide insight into ontogenetic dietary changes was examined using bottlenose dolphin tooth and skin samples. Teeth were subsampled to compare tissue produced early in life (outer tooth) to that produced later in life (inner tooth). Outer tooth had significantly higher δ¹⁵N values than the corresponding inner sample from the same tooth (n= 60, P= 0.0041), indicating that there was a temporal shift to a lower δ¹⁵N diet. There were no significant δ¹³C differences. Higher δ¹⁵N values in young have previously been attributed to the period of suckling. Analysis of skin tissue from stranded animals of different developmental stages similarly indicated that the δ¹⁵N values were significantly higher in young animals. Further comparisons indicated that the primary influence for this difference was animals with lengths less than or equal to the largest neonatal dolphin. This difference likely reflects an ontogenetic dietary shift from a sole reliance on milk to a combination of milk and prey species during the first year of life.     

Use of a serum-based glomerular filtration rate prediction equation to assess renal function by age, sex, fasting, and health status in bottlenose dolphins (Tursiops truncatus)

Glomerular filtration rate (GFR) is a direct measurement of renal function. Although clearance tests using 24‐h urine collection or blood sample series are gold standards for measuring GFR, serum‐based prediction of GFR based upon the Modification of Diet in Renal Disease (MDRD) Study equation is acceptable for routine use in human adults. The purpose of our study was to assess the ability for a modified MDRD Study equation to predict expected changes in GFR in bottlenose dolphins (Tursiops truncatus) using a healthy dolphin population represented by 1,103 routine serum samples collected from 50 dolphins of all age groups, years 1998–2005. Predicted GFR was also calculated from serum collected from a 32‐yr‐old male dolphin with end‐stage renal disease. The dolphin‐adjusted MDRD equation predicted GFR changes in our population that paralleled what has previously been reported in other mammals, including decreasing predicted GFR with age (P < 0.01), higher predicted GFR in dolphins that had recently eaten (P < 0.01), and rapidly decreasing predicted GFR in the animal with end‐stage renal disease. We conclude that a serum‐based GFR prediction equation may be a feasible means of detecting and tracking renal function in bottlenose dolphins.     

Postpartum whistle production in bottlenose dolphins

Despite much research on bottlenose dolphin signature whistles, few have investigated the role of maternal whistles in early calf development. We investigated maternal whistle use in the first weeks postpartum for captive dolphins. The overall whistling rate increased by a factor of ten when the calves were born and then decreased again in the third week of the one surviving calf. Adult whistles were distinguished from calf whistles based on the extent of frequency modulation and were further classified into signature and non-signature whistles by comparison to a dictionary of known whistles. The average rate of maternal signature whistle production increased significantly from 0.02 whistles per dolphin-minute before the calves were born to 0.2 and 0.3 whistles in weeks 1 and 2, decreasing again to 0.06 in week 3 for the mother of the surviving calf. Percent maternal signature whistles changed similarly. Signature whistle production by non-mothers did not change when the calves were born. A likely function of this increase in maternal signature whistle production is that it enables the calf to learn to identify the mother in the first weeks of life.     

Blubber cortisol qualitatively reflects circulating cortisol concentrations in bottlenose dolphins

Stress hormones, released into circulation as a consequence of disturbance, are classically assayed from blood samples but may also be detected in a variety of matrices. Blubber and fecal samples can be remotely collected from free‐ranging cetaceans without the confounding hormone elevations associated with chase, capture, and handling required to collect blood samples. The relationship between cortisol concentrations in circulation with that of blubber and feces, however, is unknown. To assess these associations, we elevated cortisol by orally administering hydrocortisone for five days in five bottlenose dolphins. Voluntary blood and fecal samples were collected daily; blubber biopsies were collected on day one, just prior to hydrocortisone administration, and days three and five of hydrocortisone administration. We evaluated subsequent changes in several circulating stress hormones as well as cortisol and glucocorticoid metabolites in blubber and feces, respectively. There was a significant association between cortisol levels in serum and in blubber (F_1,12.7 = 14.3, P < 0.01, mR² = 0.57) despite substantial variability in blubber cortisol levels. Counterintuitively, fecal cortisol metabolite levels were inversely related to serum cortisol. The relationship between serum and blubber cortisol levels suggests blubber samples from remote sampling may be useful to detect stress loads in this species.     

Brays and bits: information theory applied to acoustic communication sequences of bottlenose dolphins

Acoustic call sequences are important components of vocal repertoires for many animal species. Bottlenose dolphins (Tursiops truncatus) produce a wide variety of vocalizations, in different behavioural contexts, including some conspicuous vocal sequences – the ‘bray series’. The occurrence of brays is still insufficiently documented, contextually and geographically, and the specific functions of these multi-unit emissions are yet to be understood. Here, acoustic emissions produced by bottlenose dolphins in the Sado estuary, Portugal, were used to provide a structural characterization of the discrete elements that compose the bray series. Information theory techniques were applied to analyse bray sequences and explore the complexity of these calls. Log-frequency analysis, based on bout criterion interval, confirmed the bout structure of the bray series. A first-order Markov model revealed a distinct pattern of emission for the bray series’ elements, with uneven transitions between elements. The order in these sequential emissions was not random and consecutive decreases in higher order entropy values support the notion of a well-defined structure in the bray series. The key features of animal signal sequences here portrayed suggest the presence of relevant information content and highlight the complexity of the bottlenose dolphin’s acoustic repertoire.     

Where to catch a fish? The influence of foraging tactics on the ecology of bottlenose dolphins (Tursiops truncatus) in Florida Bay,Florida

Observations of bottlenose dolphins ( Tursiops truncatus ) in Florida Bay, Florida, between 2002 and 2005 revealed the use of three distinct foraging tactics. The goal of this study was to identify ecological correlates with tactic use and describe the impact of foraging specializations on the overall habitat use and distribution patterns of this dolphin population. Foraging tactics showed strong association with contrasting environmental characteristics, primarily depth. Locations of two of these tactic groups were spatially repulsed. Analyses of sighting histories of individual dolphins observed at foraging events determined that dolphins which employed one tactic never employed the other, and vice versa . Although bottlenose dolphins have plastic foraging behaviors, dolphins in Florida Bay appear to specialize in one tactic and subsequently limit their overall distribution patterns to coincide with habitats that facilitate success using that foraging tactic. This study demonstrates how foraging behavior can be an ecological determinant of overall dolphin habitat use patterns and works to create spatial structure within a population due to consistent mapping of tactics onto environmental variation. These foraging specializations potentially impact the social and demographic patterns of this dolphin population. The possible evolutionary mechanisms behind this intraspecific variation, including resource limitation and social learning, are considered.     

Evoked potential audiometry of 13 Pacific bottlenose dolphins (Tursiops truncatus gilli)

Evoked potential audiograms were measured in 13 Pacific bottlenose dolphins (Tursiops truncatus gilli) to determine the variability in hearing sensitivity and range of hearing. The auditory evoked potential system used a transducer embedded in a suction cup to deliver sinusoidal amplitude modulated tones to each dolphin through the pan region of the lower right jaw. Evoked potentials were recorded noninvasively using surface electrodes, and hearing thresholds were estimated by tracking the amplitude of the envelope following response, an evoked potential that is phase‐locked to the stimulus modulation rate. Frequencies tested ranged from 10 to 180 kHz in each animal. Variability in the range of hearing and age‐related reductions in hearing sensitivity and range of hearing were consistent with those observed in Atlantic bottlenose dolphins. Comparison of audiograms to a captive population of Atlantic bottlenose dolphins demonstrated that the Pacific bottlenose dolphins tested in this study had significantly lower thresholds at frequencies of 40 and 60–115 kHz. Differences in thresholds between the groups are unlikely to be due to methodological factors.     

Investigation of growth phases for bottlenose dolphins using a Bayesian modeling approach

The Gompertz function is the most commonly used growth function for cetacean studies. However, this function cannot represent multiple phases of growth. In this study, we present a Bayesian framework fitting parameters of a triple-logistic growth function to describe multiple phases of growth for bottlenose dolphins ( Tursiops truncatus ), simultaneously fitting and comparing all growth parameters between South Carolina (SC), Mississippi Sound (MSS), and Indian River Lagoon (IRL) cohorts. The fitted functions indicated a preliminary early, rapid growth phase, followed by a second phase of slower growth, and then a moderate growth spurt later in life. Growth parameters between geographic cohorts did not show obvious differences, although asymptotic length for SC dolphins was lower than MSS and IRL dolphins and significantly lower between females from SC and the IRL. Growth rate velocities between the sexes showed females exceed males initially (<1 yr), followed by males gaining an advantage around the ages of 3–4 yr until the age of around 15 yr when growth rates for both sexes approached zero (asymptotic length). This study demonstrates age-related changes in growth rates between bottlenose dolphin sexes and evidence of at least some differences ( i.e. , asymptotic length) across geographic cohorts.     

Bottlenose dolphin (Tursiops truncatus) depredation resulting in larynx strangulation with gill-net parts

Cetacean interactions with fishing gear are reported regularly and most frequently involve incidental capture. However, limited records exist related to depredation resulting in fishing gear ingestion. Here, we present cases of depredation resulting from ingestion of gill-net parts and larynx strangulation documented for the first time in a cetacean species. The bottlenose dolphin ( Tursiops truncatus ) is the only resident species in the Adriatic Sea, where small-scale commercial and private fisheries use gill nets throughout the year. A total of 120 dead-stranded bottlenose dolphins found along the Croatian coast of the Adriatic Sea were examined from 1990 to March 2008; 12 of them (10%) were affected by larynx strangulation with gill-net parts. The larynx of all affected animals showed at least one of the following pathological changes: edema, mucosal injury, and hypergranulation. In the majority of cases, it was a chronic condition. Larynx strangulations were found only in adult animals and appeared more often in animals with reduced ability to catch free-swimming prey.     

The buoyancy of the integument of Atlantic bottlenose dolphins (Tursiops truncatus): Effects of growth,reproduction, and nutritional state

In Atlantic bottlenose dolphins (Tursiops truncatus) the thickness and lipid content of blubber (the integument's specialized hypodermis) varies across ontogeny and with reproductive and nutritional state. Because the integument comprises up to 25% of total body mass in this species, ontogenetic changes in its lipid content may influence whole body buoyancy. The density and volume of the integument were measured and its buoyancy calculated across an ontogenetic series of dolphins and in pregnant and emaciated adults (total n= 45). Regional differences between the metabolically labile trunk integument and the structural tailstock integument were also investigated. Mean densities of both trunk and tailstock integument were similar across life history categories (trunk = 1,040.7 ± 14.1 kg/m³; tailstock = 1,077.1 ± 21.2 kg/m³) and were statistically similar to the density of seawater (1,026 kg/m³). The mean buoyant force of integument from the trunk (−1.01 ± 1.74 N) and tailstock (−0.30 ± 0.21 N) did not vary significantly across ontogeny. In contrast, pregnancy and emaciation did influence the integument's buoyancy, which ranged between 9 N and −45 N in these categories. Although neutral during growth, the integument's contribution to whole body buoyancy can be influenced by an individual's reproductive and nutritional status.