Recognition of classes of noise-like signals by dolphins Tursiops truncatus

The ability of the auditory system to perceive and to classify the noise-like signals imitating natural sea noises has been studied in dolphins Tursiops truncatus. Results of the studies carried out in a sea bay at free movement of the animals have shown that dolphins are able to determine noise-like signals and to ascribe them to a certain class by using invariant characters, such as rhythmical sequence of impulses, regardless of the frequency-time scale of the signal presentation.     

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.     

Blubber development in bottlenose dolphins (Tursiops truncatus)

Blubber, the lipid-rich hypodermis of cetaceans, functions in thermoregulation, buoyancy control, streamlining, metabolic energy storage, and locomotion. This study investigated the development of this specialized hypodermis in bottlenose dolphins (Tursiops truncatus) across an ontogenetic series, including fetuses, neonates, juveniles, subadults, and adults. Blubber samples were collected at the level of the mid-thorax, from robust specimens (n = 25) that stranded along the coasts of North Carolina and Virginia. Blubber was dissected from the carcass and its mass, and the depth and lipid content at the sample site, were measured. Samples were prepared using standard histological methods, viewed by light microscopy, and digital images of blubber captured. Images were analyzed through the depth of the blubber for morphological and structural features including adipocyte size, shape, and numbers, and extracellular, structural fiber densities. From fetus to adult, blubber mass and depth increased proportionally with body mass and length. Blubber lipid content increased dramatically with increasing fetal length. Adult and juvenile blubber had significantly higher blubber lipid content than fetuses, and this increase was reflected in mean adipocyte size, which increased significantly across all robust life history categories. In juvenile, subadult, and adult dolphins, this increase in cell size was not uniform across the depth of the blubber, with the largest increases observed in the middle and deep blubber regions. Through-depth counts of adipocytes were similar in all life history categories. These results suggest that blubber depth is increased during postnatal growth by increasing cell size rather than cell number. In emaciated adults (n = 2), lipid mobilization, as evidenced by a decrease in adipocyte size, was localized to the middle and deep blubber region. Thus, in terms of both lipid accumulation and depletion, the middle and deep blubber appear to be the most metabolically dynamic. The superficial blubber likely serves a structural role important in streamlining the animal. This study demonstrates that blubber is not a homogeneous tissue through its depth, and that it displays life history-dependent changes in its morphology and lipid content.     

Diaphragm muscle development in bottlenose dolphins (Tursiops truncatus)

Being born directly into the aquatic environment creates unique challenges for the breathing muscles of neonatal cetaceans. Not only must these muscles be active at the instant of birth to ventilate the lungs, but their activities must also be coordinated with those of the locomotor muscles such that breathing takes place only at the water's surface. At least one major locomotory muscle of bottlenose dolphins (Tursiops truncatus) has been demonstrated to be well developed and, therefore, able to power the neonatal dolphin's early movements (Dearolf et al. [2000] J Morphol 244:203-215). Thus, because of the demands for coordinated behavior with the locomotor muscles, it is hypothesized that the breathing muscles of bottlenose dolphins, represented in this study by the diaphragm, will also demonstrate adult morphology at birth. However, histochemical and biochemical analyses demonstrate that neonatal dolphins have immature diaphragms, with only 52% of the adult slow fiber-type profile (neonates: 34% slow-twitch fibers; adults: 66% slow-twitch fibers). The developmental state of the dolphin diaphragm is compared to those of other neonatal mammals, using a muscle development index (% slow-twitch fibers in neonatal muscle / % slow-twitch fibers in adult muscle). Fiber-type profiles reported in the literature are used to calculate index values for the diaphragms of altricial rats, rabbits, and cats, intermediate baboons and humans, and precocial sheep and horses. The dolphin is not unique in having an immature diaphragm at birth; however, there is a positive relationship between the developmental state of the diaphragm and the overall developmental state of the neonate. The presence of type IIc ("undifferentiated") fibers in the diaphragms of altricial developers (e.g., rats, rabbits, and cats) is correlated with the slow contraction speeds recorded from their diaphragms. The diaphragms of neonatal horses and dolphins express little to no type IIc fibers and, thus, may have the ability to contract at the speeds required for their increased ventilation rates. These results lead to the modification of the criterion for evaluating the developmental state of a muscle at birth. Thus, the developmental state of a neonatal muscle should be based on both its value of Dearolf et al.'s (2000) developmental index, as well as the percentage of type IIc fibers found in that muscle.     

Pulsed signal properties of free‐ranging bottlenose dolphins (Tursiops truncatus) in the central Mediterranean Sea

This study describes pulsed signals from bottlenose dolphins of the central Mediterranean Sea. Data were collected during 2011 and 2012 in 27 surveys in the Sicilian Channel, during which 163 animals were sighted. Based mainly on the pulse repetition rate, the signals were classified as Low‐frequency click (LF; single clicks without a regular pulse rate), Train click (TC; with a interclick interval of 80 ± 2 ms), Burst (with a interclick interval of 3.4 ± 0.2 ms), or Packed click (with a lower number of clicks per train and median interclick interval of 3.2 ± 0.0 ms). The measured parameters were peak sound pressure level (SPL_pk); signal duration; the 1st, 2nd, and 3rd peak of frequency; number of peaks frequency; bandwidth; centroid frequency; and the 10th, 25th, 75th, and 90th percentiles of the power spectrum distribution. Most of the parameters were significantly different among the groups, reflecting the different functions of these signals. LF clicks showed a lower peak frequency and percentiles and a longer duration and could be used to explore a wide area without a specific target focalization and with less resolution. The TC showed a higher SPL_pk, higher peak frequency, lower duration, and lower number of secondary peaks frequency, showing a better resolution to investigate a specific target.     

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.

     

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.     

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.     

Physical maturity in common bottlenose dolphins (Tursiops truncatus) from Sarasota Bay,Florida

Cetacean physical maturity is defined by growth cessation and complete fusion of epiphyses to vertebral bodies indicated by invisible sutures. Many studies have shown epiphyseal fusion is highly variable among individuals. In-depth examinations into fusion variability are lacking. We analyzed vertebrae of 37 (n = 21 female, n = 16 male) stranded common bottlenose dolphins (Tursiops truncatus) from the well-studied Gulf of Mexico, Sarasota Bay community. For each specimen, vertebrae were examined by vertebral region for degree of fusion anteriorly and posteriorly of each centrum and categorized from unfused to fused in five degrees. An ordinal logistic regression was used to estimate degree of fusion probability for each epiphysis. The model had fixed effects for age, number of offspring, sex, sexual maturity, and a random effect for epiphysis. Results show that age/reproductive status significantly explains an individual's degree of fusion. Adult females with fewer calves had more fusion than those with more reproductive experience across multiple ages. Access to long-term observational and sample data on the dolphins residing in the area served by Mote Marine Laboratory's Stranding Investigations Program offers a unique opportunity to examine the relationship between energetic demands of reproduction (calcium production/reproductive output) versus preconceived definitions of physical maturity (skeletal fusion) more closely.     

Southernmost records of bottlenose dolphins, Tursiops truncatus

Bottlenose dolphins (Tursiops spp.) are cosmopolitan animals widely distributed in waters of both hemispheres. The taxonomy of Tursiops has long been controversial, with over 20 specific names being published, and subspecies and inshore/offshore forms being proposed. In the southwestern South Atlantic, subspecies T. truncatus truncatus and T. truncatus gephyreus were proposed for specimens along the coasts of Brazil, Uruguay, and Argentina. Sightings of bottlenose dolphins are common along the coast of Argentina as far south as the Province of Chubut (ca. 46°S). Here, we summarize and discuss the southernmost records of bottlenose dolphins. We cannot make inferences about the species or subspecies to which these animals belong given the small number of specimens. Future studies of external measurements, pigmentation, DNA, and isotopes from both sides of the continent should help clarify the situation off southern South America. Furthermore, research is needed to explore a possible link between an effect of general global warming and the Tursiops specimens found this far south. The sighting and specimens described here, at 53°S–nearly 55°S, are the southernmost records for the genus and extend the range of the species in the southern South Atlantic.     

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.     

Association between burst-pulse sounds and aggressive behavior in captive Atlantic bottlenosed dolphins (Tursiops truncatus)

A specific display observed during agonistic behavior among captive Atlantic bottlenosed dolphins (Tursiops truncatus) was examined. The primary component of the display was an open-mouthed posture accompanied by violent vertical head motions and the emission of pulse-type vocalizations. Jaw-clap behavior produced during the display was quantified for use as an index of aggressive motivation. By aurally monitoring the animals' vocalizations it was found that the level of aggressive response between the participants of the interactions increased with the production and subsequent duration of burst-pulse sounds. The possibility of burstpulse sounds resulting in auditory or tactile discomfort when directed toward conspecifics is discussed.     

Association patterns of bottlenose dolphins (Tursiops truncatus) in Bahía San Antonio,Argentina

This study aims to investigate the association patterns of a small and resident population of bottlenose dolphins (Tursiops truncatus) in Bahía San Antonio (Argentina), and assess any seasonal variation in view of the reported seasonal variation in the population's grouping behavior. The estimation of social differentiation (S) was 0.29 (SE = 0.08), suggesting a homogenous society. The half‐weight index (HWI) averaged 0.23 (± 0.06), with evidence of long‐term preferred associations. However, data showed seasonal variation in the association patterns. In winter, when large groups were reported in this population, HWI values averaged 0.30 (± 0.09) with no indication of preferred or avoided associations. However, during summer, when group size was generally small, HWI values averaged 0.14 (± 0.07), with an indication of preferred/avoided associations. This study indicates that the social structure of the bottlenose dolphins in Bahía San Antonio seems relatively homogenous and flexible over time, with the formation of a random social network at times when large aggregations are formed, and a more disconnected network made up of strongly connected components when the cost of grouping is high.     

Anthropogenic injuries disrupt social associations of common bottlenose dolphins (Tursiops truncatus) in Sarasota Bay,Florida

Social connectivity is important for measuring the fitness of common bottlenose dolphins (Tursiops truncatus). While interactions in fission-fusion societies vary between individuals, studies show that repeated interactions enhance reproduction and foraging success. Injuries that potentially remove an individual from its association network may disrupt these interactions. Using data from the long-term resident dolphin community in Sarasota Bay, Florida, we investigated how anthropogenic injuries affect the dolphins' social associations by examining the differences before and after injury to individuals. We examined group size, strength, eigenvector centrality, clustering coefficient, and number of triangles and analyzed whether the animal's sex, age class, type of injury, or human intervention affected these values. We found that while group size did not change, injured dolphins had fewer preferred associates (HWI > 0.14) and were found in more fluid groups immediately after injury, but started returning to normal association levels after 2 years. This initial decrease in connectivity was not related to the age, sex, type of injury, or intervention. Despite the fluidity in individual associations, the strongest bonds remained stable, those between mothers and calves and those between male alliance partners. These findings provide some of the first information relating injuries and social networks for animals.     

Morphology of the melon and its tendinous connections to the facial muscles in bottlenose dolphins (Tursiops truncatus)

The melon is a lipid‐rich structure located in the forehead of odontocetes that functions to propagate echolocation sounds into the surrounding aquatic environment. To date, the melon's ability to guide and impedance match biosonar sounds to seawater has been attributed to its unique fatty acid composition. However, the melon is also acted upon by complex facial muscles derived from the m. maxillonasolabialis. The goal of this study was to investigate the gross morphology of the melon in bottlenose dolphins (Tursiops truncatus) and to describe how it is tendinously connected to these facial muscles. Standard gross dissection (N = 8 specimens) and serial sectioning (N = 3 specimens) techniques were used to describe the melon and to identify its connections to the surrounding muscles and blubber in three orthogonal body planes. The dolphin forehead was also thin‐sectioned in three body planes (N = 3 specimens), and polarized light was used to reveal the birefringent collagen fibers within and surrounding the melon. This study identified distinct regions of the melon that vary in shape and display locally specific muscle‐tendon morphologies. These regions include the bilaterally symmetric main body and cone and the asymmetric right and left caudal melon. This study is the first to identify that each caudal melon terminates in a lipid cup that envelopes the echolocation sound generators. Facial muscles of the melon have highly organized tendon populations that traverse the melon and insert into either the surrounding blubber, the connective tissue matrix of the nasal plug, or the connective tissue sheath surrounding the sound generators. The facial muscles and tendons also lie within multiple orthogonal body planes, which suggest that the melon is capable of complex shape change. The results of this study suggest that these muscles could function to change the frequency, beam width, and directionality of the emitted sound beam in bottlenose dolphins. The echolocation sound propagation pathway within the dolphin forehead appears to be a tunable system. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.