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.     

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.     

UNOBSERVED KILL OF NURSING DOLPHIN CALVES IN A TUNA PURSE-SEINE FISHERY

The kill of dolphins ( Stenella attenuata and S. longirostris ) in the eastern tropical Pacific tuna purse-seine fishery has been underestimated because of unobserved deaths of nursing calves due to separation from their mothers during fishing. Based on an analysis of dolphins killed from 1973 to 1990, and depending on the length at which calves are assumed to become independent, there was a deficit of calves relative to the number of lactating females killed in 24%–32% of 1,847 spotted-dolphin sets and in 13%–19% of 563 spinner-dolphin sets. We found a deficit of 0.31–0.45 spotted dolphin and 0.15–0.26 spinner dolphin calves per set. If these missing calves were added to the observed kill, it would represent an increase in the kill of 10%–15% for spotted dolphins and 6%–10% for spinner dolphins in the sets we examined. We did not attempt to estimate the actual number of unobserved calf deaths due to purse-seine fishing on dolphins, either in the sets we examined or in all dolphin sets. The actual number of unobserved calf deaths is likely to be higher than the calf deficit we found. Separation of dolphin mothers from calves could occur at any of several points in the fishing process, but most of these would be invisible to us and not produce a calf deficit. Estimation of the actual number of unobserved calf deaths would require further information on how frequently permanent mother-calf separations occur, the fraction of calves that survive after separation, the fraction of lactating females set upon that are carrying calves, and the fraction of calves killed that are actually related to lactating females killed in the same set. In any case, the observation of a calf deficit indicates that the reported dolphin kill fails to measure the full impact of purse-seine fishing on spotted and spinner dolphin populations.     

Reproduction and morphology of Central American and Tres Marias spinner dolphins in the eastern tropical Pacific

Four geographic forms of spinner dolphin (Stenella longirostris) occur in the eastern tropical Pacific. Two of the forms are pelagic: the eastern and whitebelly, and two are coastal: the Central American and Tres Marias spinner dolphins. Of these, only the Tres Marias form lacks recognition appropriate for developing management and conservation plans. Vertical aerial photographs of 57 spinner dolphin schools taken between 1988 and 2006 were used to examine the range of coastal forms and to characterize their calving seasons. On the basis of adult body length, spinner dolphins photographed off Guerrero, Mexico, were identified as those of the Central American form, which is north of their known range, and those photographed farther north, approximately 112 km off Nayarit, Mexico, were identified as the Tres Marias form. Calving peaked from October through March, and there was evidence that calving seasons differed among geographic forms with coastal forms calving later than pelagic forms. We present the first life history parameter estimates for the coastal forms of spinner dolphins, provide additional support for recognition of the Tres Marias form, and the first evidence of the Central American form occurring north of the Gulf of Tehuantepec, Mexico.     

Mitogenomic differentiation in spinner (Stenella longirostris) and pantropical spotted dolphins (S. attenuata) from the eastern tropical Pacific Ocean

Spinner dolphins (Stenella longirostris) and pantropical spotted dolphins (S. attenuata) show high intraspecific morphological diversity and endemic subspecies in the eastern tropical Pacific Ocean (ETP). Previous studies of mitochondrial DNA have found low genetic differentiation among most of these groups, possibly due to demographic factors, ongoing gene flow, and/or recent divergence. These species were heavily depleted due to bycatch in the ETP yellowfin tuna fishery. Because understanding population structure is important for accurate management of the recovery of these species, we collected whole mitochondrial genome sequences from 104 spinner and 76 spotted dolphins to test structure hypotheses at multiple hierarchical taxonomic levels. Results show differences between subspecies of spinner and spotted dolphins, but no support for the division of existing offshore stocks of spotted dolphins. We compare these results to previous results of genome‐wide nuclear SNP data and suggest high haplotype diversity, female dispersal, and/or relative power of the two data sets explains the differences observed. Interestingly, increasing the amount of mitochondrial data (base pairs and genes) did not increase ability to delimit population units. This study supports a genetic basis for management units at the subspecies level, and provides critical information for mitigating historical and continued fisheries impacts.     

TAXONOMIC STATUS AND GEOGRAPHICAL CRANIAL VARIATION OF COMMON DOLPHINS (DELPHINUS) IN THE EASTERN NORTH ATLANTIC

The common dolphin has a widespread distribution and is relatively abundant in the temperate to subtropical waters of the eastern North Atlantic. However, it is not known whether different species, subspecies, or populations occur in this region. We examined 393 common dolphin skulls obtained from both stranded and bycaught individuals collected between 1901 and 2005. The series included skulls of 152 females and 199 males, from animals ranging in body length from 93 to 230 cm and 105 to 244 cm, respectively. The ranges of total body length, skull size, RL/ZGW ratio and maximum upper alveolar (tooth) count of common dolphins in the eastern North Atlantic overlapped with those of both short- ( D. delphis ) and long-beaked ( D. capensis ) species found off the Californian coast. However, in the absence of additional data, the common dolphin in the eastern North Atlantic is regarded here as a large form of Delphinus delphis . Sexual dimorphism and possible sex-linked characters were identified within the sample. Results of the current study indicate some population differentiation within the eastern North Atlantic, with common dolphins off Portugal showing segregation in morphometric characteristics from common dolphins in other areas.     

Predictive Modeling of Spinner Dolphin (Stenella longirostris) Resting Habitat in the Main Hawaiian Islands

Predictive habitat models can provide critical information that is necessary in many conservation applications. Using Maximum Entropy modeling, we characterized habitat relationships and generated spatial predictions of spinner dolphin (Stenella longirostris) resting habitat in the main Hawaiian Islands. Spinner dolphins in Hawai'i exhibit predictable daily movements, using inshore bays as resting habitat during daylight hours and foraging in offshore waters at night. There are growing concerns regarding the effects of human activities on spinner dolphins resting in coastal areas. However, the environmental factors that define suitable resting habitat remain unclear and must be assessed and quantified in order to properly address interactions between humans and spinner dolphins. We used a series of dolphin sightings from recent surveys in the main Hawaiian Islands and a suite of environmental variables hypothesized as being important to resting habitat to model spinner dolphin resting habitat. The model performed well in predicting resting habitat and indicated that proximity to deep water foraging areas, depth, the proportion of bays with shallow depths, and rugosity were important predictors of spinner dolphin habitat. Predicted locations of suitable spinner dolphin resting habitat provided in this study indicate areas where future survey efforts should be focused and highlight potential areas of conflict with human activities. This study provides an example of a presence-only habitat model used to inform the management of a species for which patterns of habitat availability are poorly understood.     

COMPARATIVE FEEDING ECOLOGY OF SPINNER DOLPHINS (STENELLA LONGIROSTRIS) AND FRASER'S DOLPHINS (LAGENODELPHIS HOSEI) IN THE SULU SEA

Fraser's and spinner dolphins are known to feed on mesopelagic prey, bur their diets and feeding ecologies have not been compared in areas where they are observed together. In this study we examined the stomach contents of both species caught incidentally in a driftnet fishery for tuna in the eastern Sulu Sea. Importance of prey items was determined using the percent occurrence, percent number, and volumetric methods. Mesopelagic fishes, particularly myctophids (mainly Ceratoscopelus warmingi, Diaphus spp. and Myctophum asperum ), were the most important component in the diet of spinner dolphins, whereas in Eraser's dolphins, mesopelagic cephalopods ( Abraliopsis, Onychoteuthis, Histioteuthis , and Chiroteuthis ), and crustaceans ( Notostomos elegans, Acanthephyra quadrispinosa , and Acanthephyra carinata ) appeared to be equally important as the myctophid species. Fraser's dolphins appeared to feed preferentially on larger prey and had more diversified prey (Shannon's diversity index = 1.2) than spinner dolphins (diversity index = 0.9). Vertical distributions of the prey items summarized from published literature indicate that spinner dolphins forage in the upper 200 m and probably occasionally to as deep as 400 m, whereas Fraser's dolphins have a wider vertical foraging range, from near the surface to probably as deep as 600 m.     

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.     

Long‐range movement by Hector's dolphins provides potential genetic enhancement for critically endangered Maui's dolphin

For endangered populations with low genetic diversity, low levels of immigration could lead to genetic rescue, reducing the risk of inbreeding depression and enhancing chances of long‐term species survival. Our genetic monitoring of Maui's dolphins revealed the first contemporary dispersal of their sister subspecies, Hector's dolphin, from New Zealand's South Island into the Maui's dolphin distribution along ~300 km of the North Island's northwest coast. From 2010 to 2012, 44 individuals were sampled within the Maui's dolphin distribution, four of which were genetically identified as Hector's dolphins (two living females, one dead female, one dead male). We also report two Hector's dolphins (one dead female neonate, one living male) sampled along the North Island's southwest coast, outside the presumed range of either subspecies. Together, these records demonstrate long‐distance dispersal by Hector's dolphins (≥400 km) and the possibility of an unsampled Hector's dolphin population along the southwest coast of the North Island. Although two living Hector's dolphins were found in association with Maui's dolphins, there is currently no evidence of interbreeding between the subspecies. These results highlight the value of genetic monitoring for subspecies lacking distinctive physical appearances as such discoveries are not detected by other means, but have important conservation implications.     

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.     

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.     

Genetic isolation between coastal and fishery‐impacted,offshore bottlenose dolphin (Tursiops spp.) populations

The identification of species and population boundaries is important in both evolutionary and conservation biology. In recent years, new population genetic and computational methods for estimating population parameters and testing hypotheses in a quantitative manner have emerged. Using a Bayesian framework and a quantitative model‐testing approach, we evaluated the species status and genetic connectedness of bottlenose dolphin (Tursiops spp.) populations off remote northwestern Australia, with a focus on pelagic ‘offshore’ dolphins subject to incidental capture in a trawl fishery. We analysed 71 dolphin samples from three sites beyond the 50 m depth contour (the inshore boundary of the fishery) and up to 170 km offshore, including incidentally caught and free‐ranging individuals associating with trawl vessels, and 273 dolphins sampled at 12 coastal sites inshore of the 50 m depth contour and within 10 km of the coast. Results from 19 nuclear microsatellite markers showed significant population structure between dolphins from within the fishery and coastal sites, but also among dolphins from coastal sites, identifying three coastal populations. Moreover, we found no current or historic gene flow into the offshore population in the region of the fishery, indicating a complete lack of recruitment from coastal sites. Mitochondrial DNA corroborated our findings of genetic isolation between dolphins from the offshore population and coastal sites. Most offshore individuals formed a monophyletic clade with common bottlenose dolphins (T. truncatus), while all 273 individuals sampled coastally formed a well‐supported clade of Indo‐Pacific bottlenose dolphins (T. aduncus). By including a quantitative modelling approach, our study explicitly took evolutionary processes into account for informing the conservation and management of protected species. As such, it may serve as a template for other, similarly inaccessible study populations.     

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.     

SEXUAL ECOLOGY OF THE SPINNER DOLPHIN, STENELLA LONGIROSTRIS: GEOGRAPHIC VARIATION IN MATING SYSTEM

We offer the first report for a cetacean of geographical variation in mating system based in morphology. Analysis of samples from 1,678 male spinner dolphins from the eastern Pacific revealed that testis + epididymis weight was greater (to 1,354 g) in the whitebelly form of the species than in the eastern form (to 843 g). Sexual dimorphism in dorsal-fin shape is greater in the eastern form. The difference in testis size was strongly linked with shape of the dorsal fin on an individual basis. Only a few eastern males (0.6%) reached testis + epididymis weight at which all epididymides contain sperm, while a much larger proportion of whitebelly spinners (15.2%) reached this level, suggesting that a smaller proportion of eastern spinner males may participate in reproductive activity. This, and the fact that increased dimorphism and decreased testis size are indicative of increased polygyny in a wide variety of other mammal species, leads to the conclusion that the mating system varies geographically in the species, with a gradient from a more polygynous mating system in the eastern form to a more open or polygynandrous mating system in the whitebelly form. Differences in ovulation rate in the two forms are consistent with this conclusion.     

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.     

GROWTH OF BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) FROM THE INDIAN RIVER LAGOON SYSTEM, FLORIDA, U. S. A.

Gompertz growth models were fitted to total lengths and ages from tooth sections of 199 stranded bottlenose dolphins ( Tursiops truncatus ) from the Indian River Lagoon system, eastern Florida. Based on the model, dolphins from this population are estimated to be born at 119 cm and reach asymptotic length at 250 cm. No apparent pubescent growth acceleration was noted for either sex. Males appeared to grow to slightly longer lengths than females. There were small size differences between Indian River dolphins and those in Texas and Sarasota, indicating general size similarities between North Atlantic and Gulf of Mexico bottlenose dolphins stocks.     

Patterns of Dolphin Bycatch in a North-Western Australian Trawl Fishery

The bycatch of small cetaceans in commercial fisheries is a global wildlife management problem. We used data from skippers'' logbooks and independent observers to assess common bottlenose dolphin (Tursiops truncatus) bycatch patterns between 2003 and 2009 in the Pilbara Trawl Fishery, Western Australia. Both datasets indicated that dolphins were caught in all fishery areas, across all depths and throughout the year. Over the entire datasets, observer reported bycatch rates (n = 52 dolphins in 4,124 trawls, or 12.6 dolphins/1,000 trawls) were ca. double those reported by skippers (n = 180 dolphins in 27,904 trawls, or 6.5 dolphins/1,000 trawls). Generalised Linear Models based on observer data, which better explained the variation in dolphin bycatch, indicated that the most significant predictors of dolphin catch were: (1) vessel - one trawl vessel caught significantly more dolphins than three others assessed; (2) time of day – the lowest dolphin bycatch rates were between 00:00 and 05:59; and (3) whether nets included bycatch reduction devices (BRDs) - the rate was reduced by ca. 45%, from 18.8 to 10.3 dolphins/1,000 trawls, after their introduction. These results indicated that differences among vessels (or skippers'' trawling techniques) and dolphin behavior (a diurnal pattern) influenced the rates of dolphin capture; and that spatial or seasonal adjustments to trawling effort would be unlikely to significantly reduce dolphin bycatch. Recent skipper''s logbook data show that dolphin bycatch rates have not declined since those reported in 2006, when BRDs were introduced across the fishery. Modified BRDs, with top-opening escape hatches from which dolphins might escape to the surface, may be a more effective means of further reducing dolphin bycatch. The vulnerability of this dolphin population to trawling-related mortality cannot be assessed in the absence of an ongoing observer program and without information on trawler-associated dolphin community size, broader dolphin population size and connectivity with adjacent 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.