Movements and diving behavior of pelagic spotted dolphins

We radio tracked 20 pantropical spotted dolphins ( Stenella attenuata ) in the pelagic waters of the eastern tropical Pacific Ocean during three research cruises in 1992, 1993, and 2001. Dolphins were tracked for up to 6 d from 20 to 1,000 km offshore. Seventeen of these dolphins also carried time–depth recorders, nine of which were recovered with a combined total of 477 h of dive data. The movement data suggested that the dolphins associate with areas of relatively high biological productivity. Dolphins close to shore moved along the continental slope, while some dolphins traveled farther offshore along thermocline "ridges." Tracking and tagging multiple dolphins demonstrated the dynamic nature of dolphin herds, which change size and membership over the course of a day. At night, the dolphins traveled more slowly but dove deeper and longer, with more rapid ascents and descents than during daylight hours. These nighttime dive characteristics support the hypothesis suggested by some food habit studies that spotted dolphins are nocturnal feeders. Comparison of dive data with acoustic backscatter data indicates that spotted dolphins typically begin and end nighttime feeding activities with dusk and dawn diving bouts that track the vertical migration of organisms associated with the deep scattering layer.     

Why Do Dolphins Form Mixed‐Species Associations in the Azores?

Mixed‐species associations are temporary associations between individuals of different species that are often observed in birds, primates and cetaceans. They have been interpreted as a strategy to reduce predation risk, enhance foraging success and/or provide a social advantage. In the archipelago of the Azores, four species of dolphins are commonly involved in mixed‐species associations: the common dolphin, Delphinus delphis, the bottlenose dolphin, Tursiops truncatus, the striped dolphin, Stenella coeruleoalba, and the spotted dolphin, Stenella frontalis. In order to understand the reasons why dolphins associate, we analysed field data collected since 1999 by research scientists and trained observers placed onboard fishing vessels. In total, 113 mixed‐species groups were observed out of 5720 sightings. The temporal distribution, habitat (water depth, distance to the coast), behaviour (i.e. feeding, travelling, socializing), size and composition of mixed‐species groups were compared with those of single‐species groups. Results did not support the predation avoidance hypothesis and gave little support to the social advantage hypothesis. The foraging advantage hypothesis was the most convincing. However, the benefits of mixed‐species associations appeared to depend on the species. Associations were likely to be opportunistic in the larger bottlenose dolphin, while there seemed to be some evolutionary constraints favouring associations in the rarer striped dolphin. Comparison with previous studies suggests that the formation of mixed‐species groups depends on several environmental factors, and therefore may constitute an adaptive response.     

Differences in oscillatory whistles produced by spinner (Stenella longirostris) and pantropical spotted (Stenella attenuata) dolphins

Acoustic recordings of two closely related species, spinner dolphin (Stenella longirostris) and pantropical spotted dolphin (Stenella attenuata), were investigated from four different geographic locations: two in the Central Tropical Pacific, one in the Eastern Tropical Pacific and one in the Indian Ocean. The two delphinid species occur in tropical and warm temperate waters, with overlapping ranges. They produce very similar vocalizations, but at the same time their calls exhibit a certain degree of intraspecific variation among different geographic locations as has been observed in other delphinid species. Oscillatory whistles (whistles with at least two oscillations in their frequency contours) were identified and manually extracted from the recordings. Whistles with four or more maxima (oscillations) occurred only in spinner dolphins and they were present in all geographic regions investigated. In addition, the oscillatory whistles with two and three maxima were significantly more frequent in spinner than in spotted dolphins. The differences in oscillatory whistles for these two species seem to be consistent across study areas and therefore, could be used in addition to other whistle features to help distinguish between them.     

Experimental and computational biomechanical characterisation of the tracheo-bronchial tree of the bottlenose dolphin (Tursiops truncatus) during diving

Dolphins have adapted their anatomic structures to survive in the water environment and so far, the behaviour of their respiratory system during diving has not been fully understood, since they being protected species cannot be subjected to invasive analysis. Aim of this work is to model the tracheo-bronchial tree of the bottlenose dolphin Tursiops truncatus to study its behaviour during diving by coupling experimental in vitro mechanical characterisation of airways tissues to finite element computational analyses. Furthermore, a comparison was performed between the mechanical behaviour of tracheo-bronchial trees of dolphins and that of the goat, a terrestrial mammal, whose conformation of the upper airways is similar to that of the human, to determine how different structures respond to pressure in a controlled experimental set-up. The comparison between the goat and dolphin airways' mechanical behaviour highlights a lower collapsibility of the dolphin structure due to higher stiffness, lack of musculature and irregular shape of cartilaginous rings. Our data showed that the air entrapped into the airways plays a key role in avoiding the collapse. This effect is enhanced when accounting for the air flow escaping the alveoli that start to collapse during descent, even at depth as shallow as 10m of sea water. The comparison between airways behaviour of marine and terrestrial mammals may help in shedding a light on the biomechanical behaviour of human airways during breath-holding diving.     

Lung size and thoracic morphology in shallow‐ and deep‐diving cetaceans

Shallow‐diving, coastal bottlenose dolphins (Tursiops truncatus) and deep‐diving, pelagic pygmy and dwarf sperm whales (Kogia breviceps and K. sima) will experience vastly different ambient pressures at depth, which will influence the volume of air within their lungs and potentially the degree of thoracic collapse they experience. This study tested the hypotheses that lung size will be reduced and/or thoracic mobility will be enhanced in deeper divers. Lung mass (T. truncatus, n = 106; kogiids, n = 18) and lung volume (T. truncatus, n = 5; kogiids, n = 4), relative to total body mass, were compared. One T. truncatus and one K. sima were cross‐sectioned to calculate lung, thoracic vasculature, and other organ volumes. Excised thoraxes (T. truncatus, n = 3; kogiids, n = 4) were mechanically manipulated to compare changes in thoracic cavity shape and volume. Kogiid lungs were half the mass and one‐fifth the volume of those of similarly sized T. truncatus. The lungs occupied only 15% of the total thoracic cavity volume in K. sima and 37% in T. truncatus. The kogiid and dolphin thoraxes underwent similar changes in shape and volume, although the width of the thoracic inlet was relatively constrained in kogiids. A broader phylogenetic comparison demonstrated that the ratio of lung mass to total body mass in kogiids, physeterids, and ziphiids was similar to that of terrestrial mammals, while delphinids and phocoenids possessed relatively large lungs. Thus, small lung size in deep‐diving odontocetes may be a plesiomorphic character. The relatively large lung size of delphinids and phocoenids appears to be a derived condition that may permit the lung to function as a site of respiratory gas exchange throughout a dive in these rapid breathing, short‐duration, shallow divers. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Isolation and characterization of microsatellite DNA markers for spinner dolphin (<Emphasis Type="Italic">Stenella longirostris</Emphasis>)

Practically no studies on the population genetics of the spinner dolphin (Stenella longirostris) exist. Seventeen pairs of DNA primers, cloned from an Mbo I digestion of S. longirostris liver DNA, were selected from a total of 288 sequences. Eight polymorphic microsatellite DNA markers were selected from the 17 primer pairs following amplification of DNA from skin samples of 65 spinner dolphins. Characterization of the polymorphisms revealed between three and nine alleles per loci. The observed heterozygosity ranged from 0 to 0.6032, while the expected heterozygosity ranged from 0.5834 to 0.73. Seven of the eight designed primer pairs amplified DNA from three other delphinid species. There was a marked low observed heterozygosity in the spinner dolphin suggesting a high level of inbreeding within this species in the southern Atlantic.     

Striped dolphins and short‐beaked common dolphins in the Gulf of Corinth,Greece: Abundance estimates from dorsal fin photographs

Capture‐recapture methods relying on dorsal fin natural markings have never been applied successfully to striped dolphins, Stenella coeruleoalba, and were rarely used to assess abundance of short‐beaked common dolphins, Delphinus delphis. We used digital photo‐identification to obtain abundance estimates of striped and common dolphins living in mixed groups in the Gulf of Corinth, Greece. The proportion of either species was calculated based on the relative number of photographs of adult animals showing relevant portions of their body during conspicuous surfacings. Striped dolphins and common dolphins averaged 95.0% and 3.2% of all individuals, respectively. Animals showing intermediate pigmentation accounted for another 1.8%. Striped dolphin numbers were relatively high, with a point estimate of 835 animals (95% CI = 631–1,106). Common dolphins numbers were low (point estimate 28 animals; 95% CI = 11–73) and individuals were scattered within striped dolphin groups, indicating that this common dolphin population may be nonviable. Within a semiclosed Gulf exposed to considerable anthropogenic impact, the future of both dolphin species is of concern due to their suspected geographic isolation and restricted extent of occurrence. Information provided here can be used to inform timely conservation efforts.     

The dolphin cochlear nucleus: Topography,histology and functional implications

Despite the outstanding auditory capabilities of dolphins, there is only limited information available on the cytology of the auditory brain stem nuclei in these animals. Here, we investigated the cochlear nuclei (CN) of five brains of common dolphins (Delphinus delphis) and La Plata dolphins (Pontoporia blainvillei) using cell and fiber stain microslide series representing the three main anatomical planes. In general, the CN in dolphins comprise the same set of subnuclei as in other mammals. However, the volume ratio of the dorsal cochlear nucleus (DCN) in relation to the ventral cochlear nucleus (VCN) of dolphins represents a minimum among the mammals examined so far. Because, for example, in cats the DCN is necessary for reflexive orientation of the head and pinnae towards a sound source, the massive restrictions in head movability in dolphins and the absence of outer ears may be correlated with the reduction of the DCN. Moreover, the same set of main neuron types were found in the dolphin CN as in other mammals, including octopus and multipolar cells. Because the latter two types of neurons are thought to be involved in the recognition of complex sounds, including speech, we suggest that, in dolphins, they may be involved in the processing of their communication signals. Comparison of the toothed whale species studied here revealed that large spherical cells were present in the La Plata dolphin but absent in the common dolphin. These neurons are known to be engaged in the processing of low‐frequency sounds in terrestrial mammals. Accordingly, in the common dolphin, the absence of large spherical cells seems to be correlated with a shift of its auditory spectrum into the high‐frequency range above 20 kHz. The existence of large spherical cells in the VCN of the La Plata dolphin, however, is enigmatic asthis species uses frequencies around 130 kHz. J. Morphol. 2011. © 2011 Wiley Periodicals, Inc.     

A comparison of pectoral fin contact behaviour for three distinct dolphin populations

Tactile exchanges involving the pectoral fin have been documented in a variety of dolphin species. Several functions (e.g., social, hygienic) have been offered as possible explanations for when and why dolphins exchange pectoral fin contacts. In this study, we compared pectoral fin contact between dolphin dyads from three distinct dolphin populations: two groups of wild dolphins; Atlantic spotted dolphins (Stenella frontalis) from The Bahamas and Indo-Pacific bottlenose dolphins (Tursiops aduncus) from around Mikura Island, Japan; and one group of captive bottlenose dolphins (Tursiops truncatus) residing at the Roatan Institute for Marine Sciences, Anthony's Key Resort. A number of similarities were observed between the captive and wild groups, including; rates of pectoral fin contact, which dolphin initiated contact, posture preference, and same-sex rubbing partner preference. Unlike their wild counterparts, however, dolphins in the captive study group engaged in petting and rubbing at equal rates, females were more likely to contact males, males assumed the various rubbing roles more frequently than females, and calves and juveniles were more likely to be involved in pectoral fin contact exchanges. These results suggest that some aspects of pectoral fin contact behaviour might be common to many dolphin species, whereas other aspects could be species specific, or could be the result of differing environmental and social conditions.     

Morphometry and sexual dimorphism of the coastal spotted dolphin, Stenella attenuata graffmani, from Bahía de Banderas, Mexico

External measurements and size differences between the sexes were examined in the coastal spotted dolphin, Stenella attenuata graffmani , in Bahía de Banderas, on the Mexican Pacific coast. The dolphins were collected by local fishermen and 29 external characteristics were measured by members of the Marine Mammals Laboratory, University of Mexico. The length of each characteristic with respect to total length was analysed through adjustment of the data to a power equation. A stepwise discriminant analysis was applied to the absolute values and to those expressed as proportions to analyse the differences between the sexes. Results indicate that growth in these dolphins is generally negatively allometric, and most of the characteristics measured were, in both absolute and proportional terms, greater in male dolphins than in female dolphins. As found in many species of odontocetes, the discriminant analysis showed that the main differences between the sexes for this coastal subspecies include the relative positions of the umbilicus, the genital aperture and the anus. The morphometric data provided by this study, corresponding to 29 specimens of S. a. graffmani collected in a restricted locality of the Mexican Pacific coast, are particularly interesting to studies documenting latitudinal morphological differences in the coastal spotted dolphin.     

RELATIVE ABUNDANCE, DISTRIBUTION AND INTER-SPECIFIC RELATIONSHIP OF CETACEAN SCHOOLS IN THE EASTERN TROPICAL PACIFIC

The relative abundance of the most common cetacean schools in the eastern tropical Pacific Ocean for 1977–1980 are estimated based on encounter rates with tuna purse-seiners. No temporal trends were apparent in the relative abundance estimates. The geographic distributions for eight different school types are described. Multivariate statistical techniques are used to investigate interrelations between species and relationships to parameters of the physical environment. The results suggest three major species groupings: (1) an inshore grouping of bottlenose dolphins ( Tursiops truncates ), Risso's dolphin ( Grampus griseus ), pilot whales ( Globicephala macrorhynchus ) and, to a lesser extent, common dolphins ( Delphinus delphis ); (2) an offshore pelagic grouping of spotted and spinner dolphins ( Stenella attenuate and S. longirostris ); and (3) an association between pilot whales and common dolphins that overlaps the first grouping in inshore areas and also tends to be segregated from the second grouping. The results also suggest that relative densities of different school types are strongly related to physical environmental parameters, the most important being sea surface temperature, depth of the thermocline and thickness of the oxygen minimum layer.     

The development of diving bradycardia in bottlenose dolphins (<Emphasis Type="Italic">Tursiops truncatus</Emphasis>)

Bradycardia is an important component of the dive response, yet little is known about this response in immature marine mammals. To determine if diving bradycardia improves with age, cardiac patterns from trained immature and mature bottlenose dolphins (Tursiops truncatus) were recorded during three conditions (stationary respiration, voluntary breath-hold, and shallow diving). Maximum (mean: 117±1 beats·min^–1) and resting (mean: 101±5 beats·min^–1) heart rate (HR) at the water surface were similar regardless of age. All dolphins lowered HR in response to apnea; mean steady state breath-hold HR was not correlated with age. However, the ability to reduce HR while diving improved with age. Minimum and mean steady state HR during diving were highest for calves. For example, 1.5–3.5-year-old calves had significantly higher mean steady state diving HR (51±1 beats·min^–1) than 3.5–5.5-year-old juveniles (44±1 beats·min^–1). As a result, older dolphins demonstrated greater overall reductions in HR during diving. Longitudinal studies concur; the ability to reduce HR improved as individual calves matured. Thus, although newly weaned calves as young as 1.7 years exhibit elements of cardiac control, the capacity to reduce HR while diving improves with maturation up to 3.5 years postpartum. Limited ability for bradycardia may partially explain the short dive durations observed for immature marine mammals.Abbreviations ADL aerobic dive limit - cADL calculated aerobic dive limit - ECG electrocardiogram - HR heart rate - TDR time–depth recorder Communicated by L.C.-H. Wang     

Shark bite injuries on three inshore dolphin species in tropical northwestern Australia

Predation risk has a profound influence on the behavior of marine mammals, affecting grouping patterns and habitat use. Dolphins frequently bear evidence of shark bites, which can provide an indirect measure of predation pressure. Using photo‐identification data, we investigated the prevalence of shark bites on three sympatric species of inshore dolphin, the Australian snubfin (Orcaella heinsohni), Australian humpback (Sousa sahulensis), and Indo‐Pacific bottlenose dolphin (Tursiops aduncus), among four study sites in northwestern Australia. Bite prevalence varied markedly between species, with 72% of snubfin, 46% of humpback, and 18% of bottlenose dolphins exhibiting evidence of shark bites. Binomial logistic regression confirmed a high likelihood of bite presence on snubfin dolphins, and at one particular site for snubfin and bottlenose dolphins. The prevalence of tiger shark (Galeocerdo cuvier) bites on snubfin dolphins was high, and bites attributed to other carcharhinid sharks were observed on all species. While acknowledging methodological differences with other studies, the prevalence of shark bites on snubfin dolphins is among the highest reported for any dolphins, suggesting predation risk represents an important but varying influence thereon. This study provides a baseline for future investigations into the affect of predation risk on the behavioral ecology of these sympatric species.     

Thermoregulation during swimming and diving in bottlenose dolphins, Tursiops truncatus

Heat transfer from the periphery is an important thermoregulatory response in exercising mammals. However, when marine mammals submerge, peripheral vasoconstriction associated with the dive response may preclude heat dissipation at depth. To determine the effects of exercise and diving on thermoregulation in cetaceans, we measured heat flow and skin temperatures of bottlenose dolphins (Tursiops truncatus) trained to follow a boat and to dive to 15 m. The results demonstrated that skin temperatures usually remained within 1 °C of the water after all exercise levels. Heat flow from peripheral sites (dorsal fin and flukes) increased over resting values immediately after exercise at the water surface and remained elevated for up to 20 min. However, post-exercise values for heat flow from the flukes and dorsal fin decreased by 30–67% when dolphins stationed at 15 m below the surface. The pattern in heat flow was reversed during ascent. For example, mean heat flow from the flukes measured at 5 m depth, 40.10 ± 2.47 W · m⁻², increased by 103.2% upon ascent. There is some flexibility in the balance between thermal and diving responses of dolphins. During high heat loads, heat transfer may momentarily increase during submergence. However, the majority of excess heat in dolphins appears to be dissipated upon resurfacing, thereby preserving the oxygen-conserving benefits of the dive response. Accepted: 4 January 1999     

Head morphology in perinatal dolphins: a window into phylogeny and ontogeny

In this paper on the ontogenesis and evolutionary biology of odontocete cetaceans (toothed whales), we investigate the head morphology of three perinatal pantropical spotted dolphins (Stenella attenuata) with the following methods: computer-assisted tomography, magnetic resonance imaging, conventional X-ray imaging, cryo-sectioning as well as gross dissection. Comparison of these anatomical methods reveals that for a complete structural analysis, a combination of modern imaging techniques and conventional morphological methods is needed. In addition to the perinatal dolphins, we include series of microslides of fetal odontocetes (S. attenuata, common dolphin Delphinus delphis, narwhal Monodon monoceros). In contrast to other mammals, newborn cetaceans represent an extremely precocial state of development correlated to the fact that they have to swim and surface immediately after birth. Accordingly, the morphology of the perinatal dolphin head is very similar to that of the adult. Comparison with early fetal stages of dolphins shows that the ontogenetic change from the general mammalian bauplan to cetacean organization was characterized by profound morphological transformations of the relevant organ systems and roughly seems to parallel the phylogenetic transition from terrestrial ancestors to modern odontocetes.     

PHYSICAL HABITAT OF CETACEANS ALONG THE CONTINENTAL SLOPE IN THE NORTHCENTRAL AND WESTERN GULF OF MEXICO

The physical habitat of cetaceans found along the continental slope in the north-central and western Gulf of Mexico was characterized from shipboard sighting data, simultaneous hydrographic measurements, and satellite remote sensing. The study area was encompassed by the longitude of the Florida-Alabama border (87.5°W), the southernmost latitude of the Texas-Mexico border (26.0°N), and the 100-m and 2,000-m isobaths. Shipboard surveys were conducted seasonally for two years from April 1992 to May 1994. A total of 21,350 km of transect was visually sampled in an area of 154,621 km². Sighting localities of species in the study area were differentiated most clearly with bottom depth. Atlantic spotted dolphins (Stenella frontalis) were consistently found in the shallowest water on the continental shelf and along the shelf break. In addition, the bottom depth gradient (sea floor slope) was less for Atlantic spotted dolphins than for any other species. Bottlenose dolphins (Tursiops truncatus) were found most commonly along the upper slope in water significantly deeper than that for Atlantic spotted dolphins. All the other species and species categories were found over deeper bottom depths; these were Risso's dolphins (Grampus griseus), short-finned pilot whales (Glob-icephala macrorhynchus), pygmy/dwarf sperm whales (Kogia spp.), roughtoothed dolphins (Steno bredanensis), spinner dolphins (Stenella longirostris), sperm whales (Physeter macrocephalus), striped dolphins (Stenella coeruleoalba), Mesoplodon spp., pantropical spotted dolphins (Stenella attenuata), Clymene dolphins (Stenella clymene) and unidentified beaked whales (Ziphiidae). Risso's dolphins and short-finned pilot whales occurred along the upper slope and, as a subgroup, were significantly different from striped dolphins, Mesoplodon spp., pantropical spotted dolphins, Clymene dolphins, and unidentified beaked whales, which occurred in the deepest water. Pygmy/dwarf sperm whales, rough-toothed dolphins, spinner dolphins, and sperm whales occurred at intermediate depths between these two subgroups and overlapped them.     

Biphonal calls in Atlantic spotted dolphins (Stenella frontalis): bitonal and burst-pulse whistles

Biphonation, the simultaneous production of two sounds by a single animal, is found in the vocalizations of a range of mammalian species. Its prevalence suggests it plays an important role in acoustic communication. Concurrent vocal and behavioural recordings were made of Atlantic spotted dolphins (Stenella frontalis) off Bimini, The Bahamas. The occurrence of two types of biphonal signals is reported: burst-pulse whistles with combined tonal and burst-pulse elements, and bitonal whistles. Biphonal whistles are rarely described in reports of dolphin acoustic repertoires, but were common in these dolphins: of all whistles analysed (n = 1211), 26.84% were burst-pulse whistles and 4.71% were bitonal whistles. A subset of whistles (n = 397) were attributed to dolphins of specific age classes, and used to compare prevalence of biphonation across age. Biphonation occurred in 61.54% of sexually mature and 48.32% of sexually immature dolphins’ whistles. Sexually immature dolphins emitted more burst-pulse whistles than older dolphins: 44.13% of sexually immature dolphins’ whistles were burst-pulse whistles, while 15.38% of adult whistles were burst-pulse whistles. Bitonal whistle production was more prevalent in sexually mature dolphins: 41.03% of adult whistles were bitonal, while only 4.19% of sexually immature dolphins’ whistles were bitonal. The prevalence of a biphonal component in specific repeated, stereotyped whistle contours suggests that these acoustic features could be important components of contact calls, or signature whistles. The biphonal components of spotted dolphin whistles may serve to convey additional information as to identity, age or other factors to conspecifics.     

CETACEAN ABUNDANCE IN HAWAIIAN WATERS ESTIMATED FROM A SUMMER/FALL SURVEY IN 2002

Cetacean abundance is estimated for the U.S. Exclusive Economic Zone (EEZ) around the Hawaiian Islands based on a ship line‐transect survey from August to November, 2002. Sighting detection functions are estimated from this and other NOAA research surveys from 1986 to 2002 using a new, multiple‐covariate approach. Twenty‐four species were seen on this survey, including two species (Fraser's dolphin [Lagenodelphis hosei] and sei whale [Balaenoptera borealis]) that had not been previously documented to occur in Hawaiian waters. The most abundant large whales are sperm whales (Physeter macrocephalus) and Bryde's whales (Balaenoptera edeni). The most abundant delphinids are pilot whales (Globicephala macrorhynchus), rough‐toothed dolphins (Steno bredanensis), Fraser's dolphins, spotted dolphins (Stenella attenuata), and striped dolphins (Stenella coeruleoalba). Dwarf and pygmy sperm whales (Kogia sima and Kogia breviceps) and Cuvier's beaked whales (Ziphius cavirostris) are also estimated to be quite abundant. Some of the migratory baleen whales (fin whales [Balaenoptera physalus], sei whales, minke whales [B. acutorostrata], and humpback whales [Megaptera novaeangliae]) were seen only late in the survey. Abundance is estimated for 19 cetacean species. The overall density of cetaceans is low in the study area, especially for delphinids. The precision of density and abundance estimates is generally low for all species because of the low number of sightings.