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.     

RANGE CHARACTERISTICS OF PACIFIC COAST BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) IN THE SOUTHERN CALIFORNIA BIGHT

Boat-based photoidentification surveys of bottlenose dolphins (Tursiops truncatus) were conducted from 1982 to 1989 in three discrete coastal study areas within the Southern California Bight: (1) Santa Barbara, California; (2) Orange County, California; (3) Ensenada, Baja California, Mexico. A total of 207 recognizable dolphins were identified in these three “secondary” study areas. These individuals were compared to 404 dolphins identified from 1981 to 1989 in our “primary” study area, San Diego, California, to examine the coastal movement patterns of bottlenose dolphins within the Southern California Bight. A high proportion of dolphins photographed in Santa Barbara (88%), Orange County (92%), and Ensenada (88%) were also photographed in San Diego. Fifty-eight percent (n= 120) of these 207 dolphins exhibited back-and-forth movements between study areas, with no evidence of site fidelity to any particular region. Minimum range estimates were 50 and 470 km. Minimum travel-speed estimates were 11-47 km/d, and all dolphin schools sighted during the study were within 1 km of the shore. These data suggest that bottlenose dolphins within the Southern California Bight are highly mobile within a relatively narrow coastal zone. Home-range dimensions and movement patterns for many vettebrate species are influenced, in part, by variation in food resources. The unique range characteristics documented during this study may reflect the highly dynamic nature of this coastal ecosystem and the associated patchy distribution of food resources available to these bottlenose dolphins.     

How long do dolphins live? Survival rates and life expectancies for bottlenose dolphins in zoological facilities vs. wild populations

Survival rates and life expectancies are commonly agreed upon indicators of well‐being for animals in zoological facilities, but even the most recent survival statistics for bottlenose dolphins (Tursiops truncatus) in marine mammal parks and aquariums use data that are now more than 25 yr old. The current study provides a comprehensive assessment of life expectancy and survival rates for bottlenose dolphins in U.S. zoological facilities from 1974 to 2012, utilizing three different analyses (annual survival rate, age‐at‐death, and Kaplan‐Meier), examining historical trends, and comparing to comparable data from wild populations. Both survival rate and life expectancy for dolphins in zoological facilities increased significantly over the past few decades, with a modern ASR of 0.972, and mean and median life expectancies calculated via Kaplan‐Meier of 28.2 and 29.2 yr, respectively. Survival rates and life expectancies for dolphins in U.S. zoological facilities today are at least as high as those for the wild dolphin populations for which there are comparable data.     

THE DIETS OF MODERN AND HISTORIC BOTTLENOSE DOLPHIN POPULATIONS REFLECTED THROUGH STABLE ISOTOPES

The δ¹³C and δ¹⁵N compositions of teeth used in combination with existing data provide dietary information for different populations of western North Atlantic bottlenose dolphins (Tursiops truncatus). The dental isotopic signatures of bottlenose dolphins collected during the 1980s significantly differ for coastal and offshore ecotypes and are consistent with reports that coastal forms feed primarily on fish whereas offshore individuals consume more squid. In a second study, the isotopic compositions of teeth from bottlenose dolphins that span a 100-yr period and data from published stomach content analyses as well as field observations made during the past 100 yr provide evidence that coastal bottlenose dolphins from the 1880s, 1920s, and 1980s had similar diets.     

Skin lesion prevalence of estuarine common bottlenose dolphins (Tursiops truncatus) in North Carolina,with comparisons to other east coast study sites

Common bottlenose dolphins (Tursiops truncatus) are sentinels of environmental health. Skin lesions may indicate disease and can be used to infer population health. We estimated the prevalence of skin lesions and identified major lesion types on coastal bottlenose dolphins in Roanoke Sound, North Carolina, over a 3-year period using photo-identification. Boat-based surveys were conducted from April 2012 through October 2014. High quality images of distinctive fins were examined for overall prevalence (P) of any skin lesion (n = 169, P = 0.49, 95% CI = 0.42–0.57). Lesion prevalence estimates varied little between years (2012 P = 0.45, 2013 P = 0.56, 2014 P = 0.52) and most lesions were observed in the spring (P = 0.79, 95% CI: 0.57–0.92). Of six lesion types examined, pale lesions were most common (P = 0.41, 95% CI: 0.30–0.52). Annual lesion prevalence estimates for dolphins in Roanoke Sound were comparable to published estimates for T. truncatus in Charleston, South Carolina, Brunswick, Georgia, and Sarasota, Florida (p ≥ .05), although, seasonal differences in lesion occurrence and type were observed (p < .05). Future studies should examine relationships between lesions and environmental variables and use stranded dolphins to investigate skin lesion etiology.     

Using the otolith sulcus to aid in prey identification and improve estimates of prey size in diet studies of a piscivorous predator

Diet studies are fundamental for understanding trophic connections in marine ecosystems. In the southeastern US, the common bottlenose dolphin Tursiops truncatus is the predominant marine mammal in coastal waters, but its role as a top predator has received little attention. Diet studies of piscivorous predators, like bottlenose dolphins, start with assessing prey otoliths recovered from stomachs or feces, but digestive erosion hampers species identification and underestimates fish weight (FW). To compensate, FW is often estimated from the least affected otoliths and scaled to other otoliths, which also introduces bias. The sulcus, an otolith surface feature, has a species‐specific shape of its ostium and caudal extents, which is within the otolith edge for some species. We explored whether the sulcus could improve species identification and estimation of prey size using a case study of four sciaenid species targeted by fisheries and bottlenose dolphins in North Carolina. Methods were assessed first on otoliths from a reference collection (n = 421) and applied to prey otoliths (n = 5,308) recovered from 120 stomachs of dead stranded dolphins. We demonstrated in reference‐collection otoliths that cauda to sulcus length (CL:SL) could discriminate between spotted seatrout (Cynoscion nebulosus) and weakfish (Cynoscion regalis) (classification accuracy = 0.98). This method confirmed for the first time predation of spotted seatrout by bottlenose dolphins in North Carolina. Using predictive models developed from reference‐collection otoliths, we provided evidence that digestion affects otolith length more than sulcus or cauda length, making the latter better predictors. Lastly, we explored scenarios of calculating total consumed biomass across degrees of digestion. A suggested approach was for the least digested otoliths to be scaled to other otoliths iteratively from within the same stomach, month, or season as samples allow. Using the otolith sulcus helped overcome challenges of species identification and fish size estimation, indicating their potential use in other diet studies.     

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.     

Feeding aggregation and aggressive interaction between bottlenose (Tursiops truncatus) and Commerson’s dolphins (Cephalorhynchus commersonii) in Patagonia,Argentina

We report the first recorded interactions between bottlenose dolphin (Tursiops truncatus) and Commerson’s dolphins (Cephalorhynchus commersonii). The diurnal behavioral patterns of bottlenose dolphins in Bahía Engaño, Argentina, were similar to those described for other coastal populations around the world. The majority of the feeding bouts were recorded near the mouth the Chubut River. When not feeding near the river, bottlenose dolphins generally swam along the coast, and interactions with Commerson’s dolphins were recorded very close to the shore on two occasions during a 3-year period. In the first event, both species were feeding on a fish school. The second interaction was aggressive in nature, involving one juvenile and three adult bottlenose dolphins with several Commerson’s dolphins. Two of the adult bottlenose dolphins attacked the Commerson’s dolphins. We propose that the observed behavior represented defense of the juvenile bottlenose dolphin.

     

Detection of specific antibodies to morbilliviruses, Brucella and Toxoplasma in the Black Sea dolphin Tursiops truncatus ponticus and the beluga whale Delphinapterus leucas from the Sea of Okhotsk in 2002–2007

The prevalence of antibodies to morbilliviruses, Brucella and Toxoplasma was studied in the Black Sea bottlenose dolphin Tursiops truncatus ponticus and the beluga whale Delphinapterus leucas from the Sea of Okhotsk. The blood serum of 74 dolphins and 147 beluga whales was tested in 2002–2007. Antibodies to morbilliviruses were detected in 15 (20.3%) bottlenose dolphins and 20 (13.6%) beluga whales. Antibodies to Brucella were detected in 17 (23.0%) bottlenose dolphins and 10 (6.8%) beluga whales. Toxoplasma-specific antibodies were detected in 39 (52.7%) bottlenose dolphins and 7 (4.8%) beluga whales. Some animals had antibodies to two, or even three, of the pathogens. A high level of incidence of the pathogens in the sea animals was found in the densely populated coastal areas with high economic development.     

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.     

Accuracy and precision of age determination using growth layer groups for California sea lions (Zalophus californianus) with known ages

Age determination from counts of growth layer groups (GLGs) in tooth dentine is a common method for aging marine mammals. Using known‐aged animals, we validated this method for acid etched teeth of California sea lions (CSLs), Zalophus californianus. Between 1991 and 2013, the upper left canine (n = 33) was collected opportunistically during necropsy from animals tagged or branded as pups that later died. Overall, 55%–61% of age estimates by GLG counting were within 1 yr of the known‐age in the sample of 1–30‐yr‐old CSLs. Accuracy of age estimates was found to be dependent on age of the CSLs, however. 71%–79% of age estimates were within 1 yr of the known‐age in CSLs <10 yr old. These findings support the validity of counting GLGs to estimate age for CSLs <10 yr old to within 1 yr of accuracy.     

Florida red tide and brevetoxins: Association and exposure in live resident bottlenose dolphins (Tursiops truncatus) in the eastern Gulf of Mexico,U.S.A.

Bottlenose dolphins (Tursiops truncatus) along the Gulf of Mexico are frequently exposed to blooms of the toxic alga, Karenia brevis, and brevetoxins associated with these blooms have been implicated in several dolphin mortality events. Studies on brevetoxin accumulation in dolphins have typically focused on analyses of carcasses from large‐scale die‐offs; however, data are scarce for brevetoxin loads in live individuals frequently exposed to K. brevis blooms. This study investigated in vivo brevetoxin exposure in free‐ranging bottlenose dolphins resident to Sarasota Bay, Florida, utilizing samples collected during health assessments performed during multiple K. brevis blooms occurring from 2003 to 2005. Brevetoxins were detected by ELISA and LC‐MS in 63% of bottlenose dolphins sampled (n= 30) concurrently with a K. brevis bloom. Brevetoxins were present in urine and gastric samples at concentrations ranging from 2 to 9 ng PbTx‐3 eq/g, and in feces at concentrations ranging from 45 to 231 ng PbTx‐3 eq/g. Samples from individuals (n= 12) sampled during nonbloom conditions (≤1,000 cells/L) were negative for brevetoxin activity. Brevetoxin accumulation data from this study complement dolphin carcass and prey fish data from the same study area, and aid in evaluating impacts of harmful algal blooms on sentinel marine animal species along the west Florida coast.     

Age estimates derived from hard parts of swordfish Xiphias gladius from the north-western Mediterranean Sea

Accurate age estimates for fish are critical for properly understanding stock dynamics and health; this is particularly true for larger billfishes. Here we determined the most accurate aging estimation methods for swordfish (Xiphias gladius). We compared age estimates obtained from fin-ray sections, otolith sections, whole otoliths, and vertebrae collected from 87 swordfish off the east coast of Corsica. Age estimates from otolith sections were most consistently estimated across different readers (lowest average percentage error), followed by fin-ray sections, third vertebrae, and whole otoliths. When the age estimates from the otolith sections were compared with the other three age sclerochronological methods, we found the average percentage error to be lowest between the otolith section and fin-ray methods. However, age estimates from fin rays proved most useful for estimating swordfish younger than 6 years, as the fin ray-based age diverged from that of the otolith sections as the swordfish aged. Combining fin ray and otolith section techniques, we estimated the growth parameters of 1–12-year-old females (L_∞ = 259.412, k = 0.113, t₀ = −2.499) and 1–7-year-old males (L_∞ = 175.543, k = 0.202, t₀ = −2.239). We found that females grew significantly faster than males after 3 years and remained larger thereafter. Our calculated growth rates for this region of the north-western Mediterranean Sea were lower than those of the Atlantic, Pacific, and eastern Mediterranean Sea swordfish populations, and similar to growth rates recorded for the western Mediterranean Sea populations. Our study provides critical knowledge on biological-related parameters to serve as a guide for preserving the swordfish population in the Mediterranean Sea.     

Age structure of strandings and growth of Lahille's bottlenose dolphin (Tursiops truncatus gephyreus)

This study describes the age structure and sex-specific growth patterns of Lahille's bottlenose dolphins (Tursiops truncatus gephyreus), a subspecies endemic to the Southwest Atlantic Ocean (SWAO). The ages of 120 animals collected in southern Brazil between 1976 and 2017 were determined. We found high frequencies of young animals, mostly males, with no sex bias among adults. A temporal change in the age structure of strandings was observed, and we conclude it is a consequence of increased bycatch rates of young dolphins after 2000. The oldest male and female were 27 and 44 years old, respectively, suggesting that females live longer than males in southern Brazil. Growth curve analysis using Gompertz and Laird-Gompertz growth models estimated asymptotic lengths of 316.5 cm for females and 351.6 cm for males, (reached around 13 and 18 years old, respectively), supporting sexual size dimorphism for the subspecies. A second growth pulse was identified for males. Our work highlights the benefit of long-term studies and contributes valuable information toward understanding the life history of Lahille's bottlenose dolphin. It will serve as baseline for future studies that seek to understand mortality patterns of bottlenose dolphins elsewhere and the effects of anthropogenic actions on the survival of these animals.     

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.     

Genetic divergence between two phenotypically distinct bottlenose dolphin ecotypes suggests separate evolutionary trajectories

Due to their worldwide distribution and occupancy of different types of environments, bottlenose dolphins display considerable morphological variation. Despite limited understanding about the taxonomic identity of such forms and connectivity among them at global scale, coastal (or inshore) and offshore (or oceanic) ecotypes have been widely recognized in several ocean regions. In the Southwest Atlantic Ocean (SWA), however, there are scarce records of bottlenose dolphins differing in external morphology according to habitat preferences that resemble the coastal‐offshore pattern observed elsewhere. The main aim of this study was to analyze the genetic variability, and test for population structure between coastal (n = 127) and offshore (n = 45) bottlenose dolphins sampled in the SWA to assess whether their external morphological distinction is consistent with genetic differentiation. We used a combination of mtDNA control region sequences and microsatellite genotypes to infer population structure and levels of genetic diversity. Our results from both molecular marker types were congruent and revealed strong levels of structuring (microsatellites F_ST = 0.385, p < .001; mtDNA F_ST =  0.183, p < .001; Φ_ST = 0.385, p < .001) and much lower genetic diversity in the coastal than the offshore ecotype, supporting patterns found in previous studies elsewhere. Despite the opportunity for gene flow in potential “contact zones”, we found minimal current and historical connectivity between ecotypes, suggesting they are following discrete evolutionary trajectories. Based on our molecular findings, which seem to be consistent with morphological differentiations recently described for bottlenose dolphins in our study area, we recommend recognizing the offshore bottlenose dolphin ecotype as an additional Evolutionarily Significant Unit (ESU) in the SWA. Implications of these results for the conservation of bottlenose dolphins in SWA are also discussed.     

Occurrence of Indo-Pacific bottlenose dolphins (Tursiops aduncus) off the Wild Coast of South Africa using photographic identification

The present study represents the first reported boat-based photographic identification study of Indo-Pacific bottlenose dolphins (Tursiops aduncus) off the Wild Coast of southeast South Africa. This area is known for the annual sardine run, which attracts apex predators to the region during the austral winter. Dedicated photo-identification surveys were conducted along this coast at three different study sites in February, June, and November of each year from 2014 to 2016. During 47 surveys, 136 bottlenose dolphin groups were encountered, an estimated 4,474 dolphins observed, and 2,149 individuals were identified. Although most individuals (N = 1,770, 82.4%) were only observed once, some were resighted 2–7 times (N = 379, 17.6%), with an average of 305 days (range: 88–705 days) between resightings. The majority of bottlenose dolphins were resighted within the same study site (N = 192), indicating some degree of residency. However, 65 individuals were observed at two different study sites, indicating individual movements along the coast. Our findings contrast earlier suggestions that bottlenose dolphins only use the Wild Coast during the sardine run, as we found large number of animals year-round with some level of site fidelity. This highlights the importance of the Wild Coast to bottlenose dolphins and provides further information on their status off southeastern South Africa.     

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.