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Prevalence of Sarcocystis sp. in stranded Atlantic white-sided dolphins (Lagenorhynchus acutus).

In January 1998 and 1999, two mass strandings of dolphins occurred in Wellfleet, Massachusetts. The strandings were composed of 97 and 53 animals, respectively. Tissues from 35 Atlantic white-sided dolphins (Lagenorhynchus acutus) from the 1998 stranding and 52 from the 1999 stranding were examined histologically. In the 1998 stranding, unidentified protozoal tissue cysts were seen in skeletal muscle from 11 of 28 (39%) dolphins. In addition, two dolphins had a protozoal tissue cyst in cardiac muscle. In the 1999 stranding, nine of 23 (39%) dolphins had the same protozoal tissue cysts in skeletal muscle. The identification of these protozoal tissue cysts as Sarcocystis sp. was confirmed by light and transmission electron microscopy. The high prevalence of sarcocysts in these dolphins suggests that they are likely intermediate hosts for previously undescribed Sarcocystis spp. The ultrastructure of the sarcocyst walls suggests that more than one species of Sarcocystis are present in dolphins.

Animals↗

The experience of self in the bottlenose dolphin.

Marten and Psarakos have presented some evidence which suggests that objective self-awareness and possibly representations of self may characterize the dolphins' experience of self. Their research demonstrates the possibility of similarities in the sense of self between primate species and dolphins, although whether dolphins have subjective self-awareness, personal memories, and theories of self--all important facets of the sense of self in humans--was not examined. Clearly, even this limited evidence was difficult to achieve; the difficulties in adapting methods and coding behavior are quite apparent in their report. Future progress, however, may depend upon clarification of what are the necessary components for a sense of self and an explication of how these might be reflected in dolphin behavior. We are mindful of the authors' point (pp. 219 and 220) that the dolphin lives more in an acoustic than a visual environment. Thus, while tasks relying upon vision may reveal the presence or absence of the sense of self in primates, it might well be the case that in dolphins self-related experiences might be better revealed in auditory tasks. But then, what is the nature of human self-awareness in terms of audition? While both conceptual and methodological hurdles remain, Marten and Psarakos have demonstrated that important questions can be asked about the minds and phenomenal worlds of nonanthropoid species.

Animal Communication↗

A comparison of signal detection between an echolocating dolphin and an optimal receiver.

An electronic simulated target apparatus was used in a two-experiment study to compare the target detection performance of an echolocating bottlenose dolphin with an optimal receiver. Random Gaussian noise with a relatively flat spectrum from 20 to 160 kHz was used as a masking source. Experiment I was conducted to establish a technique for estimating the echo energy-to-noise ratio, Ee/N, at the dolphin's threshold of detection. Dolphins typically vary the amplitude of their emitted signal over a large range making it difficult to estimate Ee/N. In the first part of experiment I, the simulated echo was a double click, the pulses separated by 200 microseconds, with each pulse being a replica of the dolphin's transmitted signal. A staircase psychophysical procedure was used to obtain the detection threshold, and the echo energy-to-noise ratio based on the highest amplitude click emitted per trial, (Ee/N)max, was determined at each reversal point. The second echo type consisted of one of the animal's echolocation clicks, previously measured, digitized and stored in an erasable programmable read-only memory (EPROM). The electronic target simulator was modified so that every time the dolphin emitted an echolocation signal, the EPROM was triggered to produce two pulses separated by 200 microseconds. On any trial, the EPROM signal was played back at a fixed amplitude, regardless of the amplitude of the dolphin's emitted signal. The Ee/N obtained with the EPROM signal at threshold was found to be 2.9 dB lower than (Ee/N)max obtained with the normal phantom target. Therefore an estimate of Ee/N can be obtained by subtracting 2.9 dB from (Ee/N)max.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Temporal resolution of the Risso's dolphin, Grampus griseus, auditory system.

Toothed whales and dolphins (Odontocetes) are known to echolocate, producing short, broadband clicks and receiving the corresponding echoes, at extremely rapid rates. Auditory evoked potentials (AEP) and broadband click stimuli were used to determine the modulation rate transfer function (MRTF) of a neonate Risso's dolphin, Grampus griseus, thus estimating the dolphin's temporal resolution, and quantifying its physiological delay to sound stimuli. The Risso's dolphin followed sound stimuli up to 1,000 Hz with a second peak response at 500 Hz. A weighted MRTF reflected that the animal followed a broad range of rates from 100 to 1,000 Hz, but beyond 1,250 Hz the animal's hearing response was simply an onset/offset response. Similar to other mammals, the dolphin's AEP response to a single stimulus was a series of waves. The delay of the first wave, PI, was 2.76 ms and the duration of the multi-peaked response was 4.13 ms. The MRTF was similar in shape to other marine mammals except that the response delay was among the fastest measured. Results predicted that the Risso's dolphin should have the ability to follow clicks and echoes while foraging at close range.

Animal Communication↗

Laterality of flipper rubbing behaviour in wild bottlenose dolphins (Tursiops aduncus): caused by asymmetry of eye use?

To determine whether wild Indo-Pacific bottlenose dolphins (Tursiops aduncus) at Mikura Island, Japan, show asymmetry of eye or flipper use during a social behaviour, we investigated the laterality of flipper-to-body (F-B) rubbing, in which one dolphin ("rubber") rubs the body of another ("rubbee") with its flipper. We analysed 382 episodes of video-recorded F-B rubbings performed by identified individuals (N=111 rubbers). F-B rubbing was conducted significantly more frequently with the left flipper than with the right flipper. The duration of F-B rubbings was also significantly longer with the left flipper than with the right flipper. Of 20 dolphins, nine individuals showed significant left-side bias as the rubber in this behaviour, whereas no dolphins showed significant right-side bias. The results indicate a population-level left-side bias of the rubber in F-B rubbing. An analysis of the swimming configurations during this behaviour suggests that the asymmetry in F-B rubbing was caused not only by the laterality of the rubber, but by a preference for use of the left eye in both dolphins during this behaviour. Dolphins used the left eye significantly more frequently than the right eye during the inquisitive behaviour, while they showed no significant bias in flipper use during the object-carrying behaviour. These facts also suggest that the asymmetry of F-B rubbing is caused by the preference for using the left eye. Significant left-side bias was observed only in F-B rubbings initiated by the rubbee, in which the rubbee determined its position during this behaviour. This suggests that this behavioural asymmetry was enhanced by the rubbees choosing the left side of the rubber to ensure better and longer rubs.

Animals↗

Does body contact contribute towards repairing relationships? The association between flipper-rubbing and aggressive behavior in captive bottlenose dolphins.

Bottlenose dolphins exhibit complex social affiliations that may be shaped by interactions among individuals. Affiliative body contact among dolphins may repair deteriorated relationships or reduce tension within the group following aggressive interactions. We investigated the time-series association between one type of contact behavior (flipper-rubbing) and aggression by continuous observation of three captive bottlenose dolphins. For all three dolphin pairs, the elapsed time to aggressive events was significantly greater following flipper-rubbing. In two dolphin pairs comprised of a young male and an adult female, one-zero score of inter-opponent flipper-rubbing was higher for 10 min following aggressive bouts (post-AG periods) than for the same length of control (Ctrl) periods. For all three focal pairs, one-zero score of third-party rubbing was higher for post-AG than Ctrl periods. Neither the direction of rubbing nor the identity of the partner that approached prior to rubbing showed any significant tendencies. Flipper-rubbing may contribute to restore friendly relationships between former opponents or reduce conflicts in at least juvenile-adult female associations. Our results also give preliminary suggestions of the functions of third-party flipper-rubbing among bottlenose dolphins, including tension easing by the third party, or displacement as a result of aggressive interactions.

Aggression↗

Stratification and intra- and inter-specific differences in fatty acid composition of common dolphin (Delphinus sp.) blubber: implications for dietary analysis.

Sixty-five fatty acids were quantified in the blubber of common dolphins (Delphinus delphis, D. capensis) incidentally caught off the coast of southern California. Dolphins were grouped by sex, reproductive status and species, and a blubber sample was collected at a mid-lateral site located caudal to the trailing edge of the dorsal fin. Samples were divided horizontally into inner, middle and outer layers and gradients in fatty acid content (mass percent) were observed across the depth of the blubber. Levels of monounsaturated fatty acids were greatest in the outer layer, whereas levels of saturated and polyunsaturated fatty acids were greatest in the inner layer. Degree of stratification was greatest in sexually mature dolphins. Blubber of sexually immature, but physically mature, male dolphins was also highly stratified, suggesting that this difference may be attributed to differences in diet. Classification and regression tree analysis resulted in the fewest misclassifications when dolphins were grouped by species, possibly indicating that these closely related animals forage on different prey species. Dietary-derived fatty acids were typically selected as splitting criteria in classification and regression tree analyses, suggesting that the observed differences in fatty acid composition between the various groups of dolphins may be attributed to differences in diet.

Animals↗

Predominance of clockwise swimming during rest in Southern Hemisphere dolphins.

Observations on eight bottlenose dolphins located in the Southern Hemisphere during rest indicated that they spent the majority of the time (85%) engaged in behaviors that can be considered clockwise. This is in contrast with many observations of sleeping/resting dolphins in the Northern Hemisphere that spend the majority of their time involved in counterclockwise activity. This observation leads to the possibility that the reason for preferential swimming biases in dolphins is the result of global forces rather than the result of the anatomy of the individual dolphins. Our observations also indicate that dolphins change overt behavior every 40 s, coincident with the respiration rate. The possibility is suggested that the salience of neural activity controlling respiration in the reticular system may effect/disrupt reticular attentional mechanisms, thus leading to the changes in overt behavior. It is hypothesized that this 40-s period may represent the possible attention span of the sleeping bottlenose dolphin.

Activity Cycles↗

Behavioral response of Indo-Pacific humpback dolphin (Sousa chinensis) to vessel traffic.

A series of land-based surveys were conducted at two vantage points of known dolphin abundance in Hong Kong. The purpose of this study was to determine the behavioral response of Indo-Pacific humpback dolphin (Sousa chinensis) to vessel traffic. Results indicated that dolphins dove for a longer duration in areas of heavy vessel traffic or when there was the presence of an oncoming vessel. Dependent upon the type of vessel and the relative distance, dolphins might flee, continue their ongoing activity, perform a new activity, or approach the vessel. Whilst slow-moving vessels appeared not to cause immediate stress on the dolphin community, fast-moving vessels often cause disruption of behavior and social life. In order to ensure a better environment for the animals, we suggest that proactive conservation measures such as the creation of a marine park, rules and regulations for dolphin watching activities, and regional control of vessel speed should be implemented.

Animals↗

Oxygen radical generation and expression of NADPH oxidase genes in bottlenose dolphin (Tursiops truncatus) neutrophils.

Oxygen radical generation by stimulation with phorbol myristate acetate (PMA) was evaluated in bottlenose dolphin neutrophils. A Cypridina luciferin analog-dependent chemiluminescent assay demonstrated that dolphin neutrophils generate superoxide by the addition of PMA, and that its superoxide-forming activity is completely suppressed by diphenylene iodonium, a specific inhibitor of NADPH oxidase. These results indicate that dolphin neutrophils possess NADPH oxidase activity. Furthermore, the NADPH oxidase activity (hydrogen peroxide production) in dolphin neutrophils, as well as in human neutrophils, was greater at 37 degrees C than at a lower temperature. RT-PCR with specific primers revealed that dolphin neutrophils expressed the mRNAs of the major NADPH oxidase components, which included membrane-associated flavocytochrome b (gp91(phox) and p22(phox)) and cytosolic factors (p40(phox), p47(phox), and p67(phox)), implying the existence of these protein homologues in dolphin neutrophils.

Animals↗

Cloning of the IgM heavy chain of the bottlenose dolphin (Tursiops truncatus), and initial analysis of VH gene usage.

Clones encoding the dolphin IgM heavy (micro) chain gene were isolated from a cDNA library of peripheral blood leukocytes. Genomic Southern blot analyses showed that the dolphin IGHM gene is most likely present in a single copy, and its sequence shows greatest similarity to those of the IGHM gene of the sheep, pig and cow, evolutionarily related artiodactyls. The transmembrane (TM) form of the IGHM chain was isolated by 3' RACE. While showing similarities to the TM regions of other mammalian IGHM chains, the highly conserved Ser residue of the CART motif is substituted with a Gly in the dolphin. In contrast to the pig and cow, which utilize only a single VH family, the dolphin expresses at least two distinct VH families, belonging to the mammalian VH clans I and III. At least two JH genes were identified in the dolphin. Some CDR3 regions of the dolphin VH are long (up to 21 amino acids), and contain multiple Cys residues, hypothesized to stabilize the CDR3 structure through disulfide bond formation.

Amino Acid Sequence↗

Bottlenosed dolphin and human recognition of veridical and degraded video displays of an artificial gestural language.

2 bottlenosed dolphins proficient in interpreting gesture language signs viewed veridical and degraded gestures via TV without explicit training. In Exp. 1, dolphins immediately understood most gestures: Performance was high throughout degradations successively obscuring the head, torso, arms, and fingers, though deficits occurred for gestures degraded to a point-light display (PLD) of the signer's hands. In Exp. 2, humans of varying gestural fluency saw the PLD and veridical gestures from Exp. 1. Again, performance declined in the PLD condition. Though the dolphin recognized gestures as accurately as fluent humans, effects of the gesture's formational properties were not identical for humans and dolphin. Results suggest that the dolphin uses a network of semantic and gestural representations, that bottom-up processing predominates when the dolphin's short-term memory is taxed, and that recognition is affected by variables germane to grammatical category, short-term memory, and visual perception.

Animal Communication↗

Travel at low energetic cost by swimming and wave-riding bottlenose dolphins.

Over the past 50 years there has been much speculation about the energetic cost of swimming and wave-riding by dolphins. When aligned properly in front of the bow of moving ships in the stern wake of small boats, on wind waves, and even in the wake of larger cetaceans, the animals appear to move effortlessly through the water without the benefit of propulsive strokes by the flukes. Theoretically, body streamlining as well as other anatomical and behavioural adaptations contribute to low transport costs in these animals. The economy of movement permitted by wave-riding has been perceived as an energetic advantage for the swimming dolphin, but has been hard to prove in the absence of physiological data for exercising cetaceans. Here we determine the aerobic and anaerobic costs of swimming and wave-riding in bottlenose dolphins and find that the minimum cost of transport for swimming dolphins is 1.29 +/- 0.05 J kg-1 m-1 at a cruising speed of 2.1 m s-1. Aerobic costs are nearly twice as high for swimming seals and sea lions, and 8-12 times higher for human swimmers. Wave-riding by dolphins provides additional benefits in terms of speed. The results indicate that behavioural, physiological and morphological factors make swimming an economical form of high-speed travel for dolphins.

Animals↗

Fine-scale behaviour of bottlenose dolphins around gillnets.

We studied the fine-scale behaviour of bottlenose dolphins Tursiops truncatus around gillnets in North Carolina, USA, during May and June 2002. We made observations from an overhead digital video camera, suspended from a helium-filled aerostat, tethered 70 m above a fishing vessel. We positioned the camera above a gillnet set for Spanish mackerel Scomberomorus maculatus. We observed frequent encounters (n = 36) and interactions (n = 27) between dolphins and the net, but no dolphins became entangled. Most dolphins diverted their course around the net, but on nine occasions we observed dolphins engaging in depredation. We conclude that interactions between dolphins and these gillnets are common, but that entanglement is rare.

Animals↗

Evolution of river dolphins.

The world's river dolphins (Inia, Pontoporia, Lipotes and Platanista) are among the least known and most endangered of all cetaceans. The four extant genera inhabit geographically disjunct river systems and exhibit highly modified morphologies, leading many cetologists to regard river dolphins as an unnatural group. Numerous arrangements have been proposed for their phylogenetic relationships to one another and to other odontocete cetaceans. These alternative views strongly affect the biogeographical and evolutionary implications raised by the important, although limited, fossil record of river dolphins. We present a hypothesis of river dolphin relationships based on phylogenetic analysis of three mitochondrial genes for 29 cetacean species, concluding that the four genera represent three separate, ancient branches in odontocete evolution. Our molecular phylogeny corresponds well with the first fossil appearances of the primary lineages of modern odontocetes. Integrating relevant events in Tertiary palaeoceanography, we develop a scenario for river dolphin evolution during the globally high sea levels of the Middle Miocene. We suggest that ancestors of the four extant river dolphin lineages colonized the shallow epicontintental seas that inundated the Amazon, Paraná, Yangtze and Indo-Gangetic river basins, subsequently remaining in these extensive waterways during their transition to freshwater with the Late Neogene trend of sea-level lowering.

Animals↗

Signature-whistle production in undisturbed free-ranging bottlenose dolphins (Tursiops truncatus).

Data from behavioural observations and acoustic recordings of free-ranging bottlenose dolphins (Tursiops truncatus) were analysed to determine whether signature whistles are produced by wild undisturbed dolphins, and how whistle production varies with activity and group size. The study animals were part of a resident community of bottlenose dolphins near Sarasota, Florida, USA. This community of dolphins provides a unique opportunity for the study of signature-whistle production, since most animals have been recorded during capture-release events since 1975. Three mother-calf pairs and their associates were recorded for a total of 141.25 h between May and August of 1994 and 1995. Whistles of undisturbed dolphins were compared with those recorded from the same individuals during capture-release events. Whistles were conservatively classified into one of four categories: signature, probable signature, upsweep or other. For statistical analyses, signature and probable signature whistles were combined into a 'signature' category; upsweep and other whistles were combined into a 'non-signature' category. Both 'signature' and 'non-signature' whistle frequencies significantly increased as group size increased. There were significant differences in whistle frequencies across activity types: both 'signature' and 'non-signature' whistles were most likely to occur during socializing and least likely to occur during travelling. There were no significant interactions between group size and activity type. Signature and probable signature whistles made up ca. 52% of all whistles produced by these free-ranging bottlenose dolphins.

Animals↗

Evidence for male dispersal along the coasts but no migration in pelagic waters in dusky dolphins (Lagenorhynchus obscurus).

Using nine nuclear species-specific microsatellite loci and two mitochondrial gene fragments (cytochrome b and control region), we investigated the processes that have shaped the geographical distribution of genetic diversity exhibited by contemporary dusky dolphin (Lagenorhynchus obscurus) populations. A total of 221 individuals from four locations (Peru, Argentina, southern Africa, and New Zealand) were assayed, covering most of the species' distribution range. Although our analyses identify a general demographic decline in the Peruvian dusky dolphin stock (recently affected by high natural and human-induced mortality levels), comparison between the different molecular markers hint at an ancient bottleneck that predates recent El Niño oscillations and human exploitation. Moreover, we find evidence of a difference in dispersal behaviour of dusky dolphins along the South American coast and across the Atlantic. While data in Peruvian and Argentine waters are best explained by male-specific gene flow between these two populations, our analyses suggest that dusky dolphins from Argentina and southern Africa recently separated from an ancestral Atlantic population and, since then, diverged without considerable gene flow. The inclusion of a few New Zealand samples further confirms the low levels of genetic differentiation among most dusky dolphin populations. Only the Peruvian dusky dolphin stock is highly differentiated, especially at mitochondrial loci, suggesting that major fluctuations in its population size have led to an increased rate of genetic drift.

Africa, Southern↗

Echolocation in the Risso's dolphin, Grampus griseus.

The Risso's dolphin (Grampus griseus) is an exclusively cephalopod-consuming delphinid with a distinctive vertical indentation along its forehead. To investigate whether or not the species echolocates, a female Risso's dolphin was trained to discriminate an aluminum cylinder from a nylon sphere (experiment 1) or an aluminum sphere (experiment 2) while wearing eyecups and free swimming in an open-water pen in Kaneohe Bay, Hawaii. The dolphin completed the task with little difficulty despite being blindfolded. Clicks emitted by the dolphin were acquired at average amplitudes of 192.6 dB re 1 microPa, with estimated sources levels up to 216 dB re 1 microPa-1 m. Clicks were acquired with peak frequencies as high as 104.7 kHz (Mf(p) = 47.9 kHz), center frequencies as high as 85.7 kHz (Mf(0) = 56.5 kHz), 3-dB bandwidths up to 94.1 kHz (M(BW) = 39.7 kHz), and root-mean-square bandwidths up to 32.8 kHz (M(RMS) = 23.3 kHz). Click durations were between 40 and 70 micros. The data establish that the Risso's dolphin echolocates, and that, aside from slightly lower amplitudes and frequencies, the clicks emitted by the dolphin were similar to those emitted by other echolocating odontocetes. The particular acoustic and behavioral findings in the study are discussed with respect to the possible direction of the sonar transmission beam of the species.

Animals↗