PHYSIOLOGICAL AND BEHAVIORAL DEVELOPMENT IN DELPHINID CALVES: IMPLICATIONS FOR CALF SEPARATION AND MORTALITY DUE TO TUNA PURSE-SEINE SETS

Tuna purse-seiners in the eastern tropical Pacific (ETP) capture yellowfin tuna by chasing and encircling herds of associated dolphins. This fishery has caused mortality in 14 dolphin species (20 stocks) and has led to significant depletions of at least three stocks. Although observed dolphin mortality is currently low, set frequency remains high and dolphin stocks are not recovering at expected rates. Mortality of nursing calves permanently separated from their mothers during fishery operations may be an important factor in the lack of population recovery, based on the recent discovery that calves do not accompany 75%–95% of lactating females killed in the purse-seine nets. We assessed age-specific potential for mother–calf separations and subsequent mortality of calves by reviewing and synthesizing published data on physiological and behavioral development in delphinids from birth through 3 yr postpartum. Results indicate that evasive behavior of mothers, coupled with the developmental state of calves, provides a plausible mechanism for set-related mother–calf separations and subsequent mortality of calves. Potential for set-related separation and subsequent mortality is highest for 0–12-mo-old dolphins and becomes progressively lower with age as immature dolphins approach adult stamina and attain independence.     

AGE AND LENGTH AT WEANING AND DEVELOPMENT OF DIET OF PANTROPICAL SPOTTED DOLPHINS, STENELLA ATTENUATA, FROM THE EASTERN TROPICAL PACIFIC

Using stomach contents from 203 spotted dolphins ( Stenella attenuata ) killed in the yellowfin tuna fishery, we modeled the weaning process of calves. Spotted dolphins began to take solid food at approximately 6 mo of age, or 115 cm, but continued to suckle until they were nearly 2 yr old. Calves tended to feed more frequently on squid as they got older, which suggested there was a shift in diet during weaning. The average age and total body length at weaning was estimated to be 0.8 yr (approximately 9 mo) and 122 cm. The oldest suckling calf was almost 2 yr old, which suggests that some calves continued to suckle for more than a year after they could have been weaned. A better understanding of the weaning process, especially quantifying the period of time when calves are nutritionally dependent on their mothers may lead to a better evaluation of their potential vulnerability to the disturbance caused by the yellowfin tuna purse-seine fishery.     

SPOTTED DOLPHIN EVASIVE RESPONSE IN RELATION TO FISHING EFFORT

Spotted dolphins in the eastern Pacific Ocean associate with yellowfin tuna. During the chase and encirclement phases of purse-seining for tunas, dolphin attempt to evade encirclement with the purse-seine net. We used data on evasive behavior (1982–2001) and numbers of purse-seine sets (1959–2001) to study the relationship between evasion and fishing effort. Results show that in nearshore areas first exploited by the fishery in the early 1960s, dolphins exhibited high evasion, but with a limited correlation between evasion and cumulative effort. In areas farther offshore next exploited in the mid-to late-1960s, dolphins showed high evasion and a significant correlation between evasion and cumulative effort. Dolphins in far-western and southern areas, first exploited in the late 1960s to early 1970s, exhibited low evasion, with little relationship to cumulative effort. We hypothesize that this spatial pattern is the result of two types of pressure from fishing: early effort in nearshore areas with a high risk of mortality that generated a lasting evasive response, followed by a longer period of even greater effort but with lower risk of mortality that generated evasion by longer-term learning.     

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.     

Hearing loss in stranded odontocete dolphins and whales

The causes of dolphin and whale stranding can often be difficult to determine. Because toothed whales rely on echolocation for orientation and feeding, hearing deficits could lead to stranding. We report on the results of auditory evoked potential measurements from eight species of odontocete cetaceans that were found stranded or severely entangled in fishing gear during the period 2004 through 2009. Approximately 57% of the bottlenose dolphins and 36% of the rough-toothed dolphins had significant hearing deficits with a reduction in sensitivity equivalent to severe (70-90 dB) or profound (>90 dB) hearing loss in humans. The only stranded short-finned pilot whale examined had profound hearing loss. No impairments were detected in seven Risso's dolphins from three different stranding events, two pygmy killer whales, one Atlantic spotted dolphin, one spinner dolphin, or a juvenile Gervais' beaked whale. Hearing impairment could play a significant role in some cetacean stranding events, and the hearing of all cetaceans in rehabilitation should be tested.     

THE LIFE HISTORY OF FREE-RANGING ATLANTIC SPOTTED DOLPHINS (STENELLA FRONTALIS): AGE CLASSES, COLOR PHASES, AND FEMALE REPRODUCTION

Atlantic spotted dolphins (Stenella frontalis) were observed underwater and from the surface from 1985 to 1996 and photographed through successive years. Individuals were categorized into age classes by their degree of spotting and color phases. Dolphins spent an average of 3 yr in the two-tone color phase, 5 yr in the speckled phase, 7 yr in the mottled phase and up to 10 yr or more in the fused phase.
Sex ratios were close to parity, with old adults skewed towards females and juveniles and young adults skewed towards males. The average calving interval for 24 females was 2.96 years with a range of 1–5 yr. Females whose calves survived the first year had a significantly longer calving interval (3.56 years). The ages of first parturition for five females were estimated to be 10–12 yr. The age at sexual maturation was estimated to range from 8 to 15 yr.
Pregnancy rate fluctuated annually, with an average rate of 0.25 (range 0.07–0.57). Annual average birth rate was 0.08 (range 0.07–0.14), average calf production was 0.33 (range 0.06–0.52), average fecundity was 0.23 (range 0.13–0.30), and average recruitment was 0.06 (range 0.03–0.08). Most females who lost a calf conceived the same or following year.
Lactation lasted up to 5 yr, and 45% of visibly pregnant females were also lactating. Age of first parturition was associated with the mottled color phase. Average first-year mortality rate of calves was 0.24.     

Evidence for infanticide in bottlenose dolphins: an explanation for violent interactions with harbour porpoises?

Most harbour porpoises found dead on the north-east coast of Scotland show signs of attack by sympatric bottlenose dolphins, but the reason(s) for these violent interactions remain(s) unclear. Post-mortem examinations of stranded bottlenose dolphins indicate that five out of eight young calves from this same area were also killed by bottlenose dolphins. These data, together with direct observations of an aggressive interaction between an adult bottlenose dolphin and a dead bottlenose dolphin calf, provide strong evidence for infanticide in this population. The similarity in the size range of harbour porpoises and dolphin calves that showed signs of attack by bottlenose dolphins suggests that previously reported interspecific interactions could be related to this infanticidal behaviour. These findings appear to provide the first evidence of infanticide in cetaceans (whales, dolphins and porpoises). We suggest that infanticide must be considered as a factor shaping sociality in this and other species of cetaceans, and may have serious consequences for the viability of small populations.     

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.     

Long‐term social structure of a resident community of Atlantic spotted dolphins,Stenella Frontalis,in the Bahamas 1991–2002

Long‐term social structure data on small delphinids is lacking for most species except the bottlenose dolphin. This study describes the long‐term social structure of one community of Atlantic spotted dolphins, Stenella frontalis, divided into three social clusters. Data from 12 yr were analyzed using SOCPROG 2.3. Coefficients of association (CoA) were calculated using the half‐weight index. The overall mean community CoA ranged from 0.09 to 0.12. Temporal analyses and mantel tests revealed significant differences between sex class associations due to high male‐male CoA (0.12–0.23) compared to female‐female and mixed sex CoA (0.08–0.10). Female associations were strongly influenced by reproductive status, calf care, and social familiarity, but not by age class. Male associations were strongly influenced by age, access to females, and alliance formation. Males showed two levels of alliance formation, long‐term first order pairs/trios (CoA 0.70–1.00) and shorter‐term second order alliances between two or more first order alliances (CoA 0.45–0.69), and a possible third level during interspecies interactions. Mating strategies, sex, and cluster formation shaped the social structure in this spotted dolphin community. Similar to many bottlenose dolphin studies, long‐term affiliations for spotted dolphins were correlated with age, sex, and reproductive status.     

A DWARF FORM OF THE SPINNER DOLPHIN (STENELLA LONGIROSTRIS) FROM THAILAND

A very small form of the spinner dolphin has been found to inhabit the Gulf of Thailand. Ten specimens taken incidentally in a local shrimp fishery differ from specimens of this species collected elsewhere in body size and shape, skull size and shape, number of teeth and numbers of thoracic and lumbar vertebrae. Four cranially adult males were 129–137 cm long, well below the ranges for the Indian Ocean, western Pacific, central Pacific, eastern Pacific and Atlantic. The skull is also very small. Tooth counts and vertebral counts average lower than in other series. The color pattern is not significantly different from that of spinner dolphins in the central and western Pacific, Atlantic and Indian Oceans but differs from that of the small eastern spinner of the eastern Pacific. The Gulf of Thailand specimens are morphologically separable from all other specimens, but it is to be expected that when larger samples are available there will be some overlap. The dwarf form may overlap in body and skull size with small spinner dolphins taken incidentally in a gillnet fishery for sharks off northern Australia. The geographic range of the dwarf spinner may be restricted. The size and status of the population and the impact of the shrimp fishery are unknown and should be investigated. The dwarf spinner may have an ecology different from that of other spinners, feeding mainly on reef-associated and benthic organisms rather than mesopelagic animals.     

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.     

DIEL PATTERNS IN AGGREGATIONS OF PELAGIC DOLPHINS AND TUNAS IN THE EASTERN PACIFIC

Data collected by scientific technicians aboard tuna purse seiners in the eastern Pacific Ocean since the early 1970s have allowed us to study the biology and herd dynamics of pelagic dolphins. A pattern of increasing group size in the morning and subsequent decline in the late afternoon or night was evident for spotted, spinner, and common dolphins, as well as for large yellowfin tuna that associate with dolphins. Diel patterns were also apparent in the formation of mixed-species herds of spotted and spinner dolphins and tuna-dolphin aggregations. Different patterns, however, were displayed by bottlenose dolphins and by yellowfin and skipjack tunas that did not associate with dolphins. It appears that these diel patterns are produced by an interaction of predation pressure and prey distribution.     

Studying small purse‐seine vessel fishing behavior with tuna catch data: Implications for eastern Pacific Ocean dolphin conservation

Despite achievements in dolphin conservation for the tuna purse‐seine fishery of the eastern Pacific Ocean, debate continues about the magnitude and importance of dolphin mortality caused by small (unobserved) vessels. In‐port sampling of tuna catch size composition is a potentially cost‐effective means of identifying unobserved vessels that may be catching tunas associated with dolphins because yellowfin tuna caught in association with dolphins are larger, on average, than those caught in other types of purse‐seine sets. A classification algorithm to predict purse‐seine set type (“dolphin” vs. “nondolphin”) was built from port‐sampling data on yellowfin tuna length‐frequencies and the date and location of fishing of large (observed) vessels. This classification algorithm was used to screen the port‐sampling data of small vessels collected during 2006‐2009, assuming the fishing practices of the two groups resulted in similar catch characteristics. From these results, hypothetical time series of dolphin mortality for small vessels were constructed and incorporated into a population dynamics model, along with mortalities of large vessels. Results suggest that any dolphin mortality of small vessels is unlikely to be substantially affecting trends in dolphin abundance. These results underscore the importance of in‐port sampling, in combination with at‐sea observation and fishery‐independent surveys, to effective management.     

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.     

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.     

SPINNER DOLPHINS (STENELLA LONGIROSTRIS) OF THE WESTERN PACIFIC AND SOUTHEAST ASIA: PELAGIC AND SHALLOW-WATER FORMS1

A dwarf form of the spinner dolphin has been reported from the Gulf of Thailand, while more typical large spinner dolphins have been described from Japanese waters and other localities in the western Pacific. These reports have been based on very few specimens. Our purpose in this study was to determine the affinities of spinner dolphins throughout the region based on larger samples and to review their taxonomic status, with an hypothesis of two widespread ecotypic forms, or subspecies. We examined 213 osteological specimens, from a tuna gillnet fishery in the Philippines, from the former Taiwanese shark gillnet fishery in the Timor and Arafura Seas off northern Australia, from the Gulf of Thailand, from other areas in the western Pacific and Southeast Asia, from the eastern Indian Ocean, and from the Central and South Pacific. Results show that spinner dolphins from the deep inner waters of the Philippines conform to the large pelagic type of spinner dolphin that inhabits the Central and South Pacific, the western Pacific and the eastern Indian Ocean. The skull is similar in size and shape to the holotype specimen of S. longirostris (from unknown locality). This form feeds primarily on small mesopelagic fishes and squids. Spinner dolphins from the shallow waters of inner Southeast Asia represented in the sample, including the Gulf of Thailand, Timor Sea and Arafura Sea, are smaller in body and skull size, have fewer teeth and vertebrae, and feed mainly on benthic and coral reef fishes and invertebrates. We hypothesize that this form also inhabits the Java Sea and other shallow waters throughout inner Indonesia and Malaysia. We redescribe a subspecies corresponding to the small form and based on Delphinus roseiventris Wagner 1846 from the Arafura Sea, designating a neotype and paraneotype specimens.     

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.     

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.     

Circadian and seasonal rhythms in the behavior of spinner dolphins (Stenella longirostris)

The present study investigated both circadian and seasonal fluctuations in the daytime activities of the spinner dolphin, Stenella longirostris , from the Fernando de Noronha Archipelago in Brazil. The number of dolphins, and aerial, and reproductive activities were documented. The observations were carried out from January 1997 to December 2001. Temporal series and rhythmic characteristics (mesor, rhythmic percentage, and acrophase) were obtained by COSINOR analysis and later compared. The dolphins entered the bay in the morning, displayed aerial and reproductive activities during daytime, and left the bay in late afternoon to the open ocean. This study indicated that the rainy season affected the three behaviors investigated decreasing the rhythms parameter and advancing the beginning of these activities. The number of individuals was higher during the dry season and the animals stayed longer inside the bay. During the dry season, there was a bimodal expression of aerial activity, expressing a longer use of the temporal niche than in the rainy season. The phases with high frequencies of aerial activity seemed associated with those showing high reproductive activity, both with peak frequencies at about 0800. The results represent an important contribution to the advancement of chronobiological studies, and to the biology of cetaceans, considering that the existence of circadian and seasonal rhythms was proven in the behavior of spinner dolphins in an area of the SW Atlantic. Moreover, it allows restricting periods of the day for the activities of tourism as a form of minimizing the impacts of the boats on the dolphins.