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.     

GEOGRAPHIC VARIATION IN TOOTH MORPHOLOGY AND DENTINAL PATTERNS IN THE SPINNER DOLPHIN, STENELLA LONGIROSTRIS

Sixteen tooth characters were analyzed in 277 specimens of spinner dolphins ( Stenella longirostris ) from the eastern tropical Pacific to determine whether the characters were associated with size of the animal, species-stock membership or geographic location. The tooth characters included 5 continuous measurements and 11 qualitative variables. Statistical analyses of the characters included two-way ANOVA tests which assessed continuous measurement variables for differences based on stock membership, latitude and longitude of collection or whether the animal was collected north or south of the equator. Principal component analyses combined factors of continuous variables with those of qualitative variables into components which were used as variables in further analyses. Discriminant analyses were performed using continuous variables alone and using principal components as potentially discriminating variables. Results showed that tooth differences were not associated with size but significant differences were found in some tooth variables with stock membership, incremental changes in latitude and whether the animal was collected north or south of the equator. A discriminant function, containing only tooth length, correctly classified 67% of northern and 78% of southern animals according to location of collection north or south of the equator.     

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.     

LIFE HISTORY OF FRASER'S DOLPHIN, LAGENODELPHIS HOSEI, BASED ON A SCHOOL CAPTURED OFF THE PACIFIC COAST OF JAPAN

Life history of Fraser's dolphin, a little known delphinid species, was examined based on 108 specimens from a school captured by the driving fishing method in Japan. The sex ratio was approximately 1:1, and mature dolphins of both sexes formed the bulk of the school. The oldest animals were two males and a female of 17.5 yr. Age and body length at sexual maturity were estimated at 7–10 yr and 220–230 cm in males and 5–8 yr and 210–220 cm in females. Mature males were larger in body length than mature females and showed apparent secondary sexual features: deepening of the tail stock and widening and darkening of the lateral dark stripe. The annual ovulation rate was 0.49. The estimated neonatal length (110 cm) predicts a gestation period of about 12.5 mo and calving peaks in spring and probably also in fall. The calving interval was estimated to be about 2 yr. These life history parameters are similar to those of the striped and pantropical spotted dolphins, except for longevity. The reproductive rate of this species may be lower than that of other pelagic delphinids, if the observed shorter longevity is real.     

CRANIAL SEXUAL DIMORPHISM AND GEOGRAPHIC VARIATION IN ERASER'S DOLPHIN, LAGENODELPHIS HOSEI

Knowledge of geographic variation is important to questions of population assessment and management. Fraser's dolphins have been exploited in two regions in the western Pacific. Analysis of 137 skulls from the Philippines, Japan, Taiwan, Australia, Indonesia, Malaysia, South Africa, France, the U.S., St. Vincent and the Grenadines, and the eastern tropical Pacific revealed sexual dimorphism in 5 of 26 measurements (difference of 1.9%-5.8% between males and females), similar to levels of cranial dimorphism in other small pelagic delphinids. Males had a larger braincase and temporal fossae and smaller external nares than females. Sexually dimorphic characters were excluded, and male and females samples were pooled to examine geographic differences in the remaining characters. Multivariate analyses yielded significant differences between the Philippine and Japanese series within the North Pacific and between a pooled North Pacific series and a North Atlantic series. The Japanese skulls were on average broader and had a wider rostrum, larger orbit, larger internal nares, and longer braincase than the Philippine skulls. These differences suggest that Fraser's dolphins exploited in Japanese and Philippine waters in directed fisheries or as by catch belong to different populations and should be assessed and managed separately.     

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.     

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.     

条纹原海豚mtDNA控制区序列变异分析

条纹原海豚( Stenella coeruleualba )是一种暖水性的小型鲸类,在50°N～40°S之间有分布,在其分布区内的许多水域,均受到渔业误捕或直接捕捞的影响(Jefferson et al .,1993).但由于缺乏该物种全面而系统的种群生物学知识,IUCN红皮目录将其保护生物学地位定为"不清楚"(Hilton-Taylor,2000),需要进一步开展保护生物学研究.     

MITOCHONDRIAL DNA VARIABILITY OF STRIPED DOLPHINS (STENELLA COERULEOALBA) IN THE SPANISH MEDITERRANEAN WATERS

Frozen muscle samples from 44 striped dolphins stranded on the Spanish Mediterranean coasts from 1990 to 1993 have been studied by means of mitochondrial DNA (mtDNA) restriction site analysis. Thirty-five of these dolphins were affected by a die-off occurring during this time in the western Mediterranean Sea. The mtDNA from each dolphin was digested with 15 restriction endonucleases that recognized 61 different restriction sites. The specific location of these sites on the mitochondrial gene map allowed us to determine the distribution of variability along this molecule. From the restriction analysis, a total of 15 different composite patterns or haplotypes was obtained and their phylogenetic relationships established both by cladistic and phenetic methods.
Estimates of sequence divergence among the 15 haplotypes ranged from 0.148% to 0.919%. Haplotype diversity (average heterozygosity) obtained was 0.789, a high value compared to other available data on cetaceans, especially considering that only a small area of the distribution of the species has been sampled. Neither population subdivision nor any evidence of correlation between haplotype frequencies and temporal or geographic distribution of dolphins has been detected.     

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.     

REPRODUCTIVE MATURITY AND SEASONALITY OF MALE SPOTTED DOLPHINS, STENELLA ATTENUATA, IN THE EASTERN TROPICAL PACIFIC

We estimated age at attainment of sexual maturity and examined reproductive seasonality for male spotted dolphins, Stenella attenuata , from the eastern tropical Pacific Ocean. Maturity was determined by histological examination of testes. Average age at sexual maturation was 14.7 yr (the mean of two readers' age estimates). Testis and epididymis weight and seminiferous tubule diameters were reliable indicators of maturity, whereas age, length and color phase were less reliable. Seasonality was determined by changes in testis and epididymis weight, relative quantity of spermatids and spermatozoa, and lumen diameter, as well as an index of testis development (weight of the right testis and epididymis divided by length of the right testis). Testis and epididymis weights and index values peaked in July and August, midway between two predicted mating seasons for the northern offshore stock, but spermatozoa levels were elevated during the predicted breeding seasons.     

GEOGRAPHIC PATTERNS OF VARIATION IN OFFSHORE SPOTTED DOLPHINS (STENELLA ATTENUATA) OF THE EASTERN TROPICAL PACIFIC OCEAN

The spotted dolphin (Stenella attenuata), a species commonly associated with the tuna fishery, is widely distributed in the eastern tropical Pacific Ocean (ETP). Geographic patterns in morphological characteristics of this dolphin were evaluated for 36 skull measurements of 574 museum specimens allocated to 5° latitude-longitude blocks. A strong subdivision between northern and southern offshore populations was demonstrated using principal components, canonical variates and cluster (UPGMA and function-point) analyses of block means derived from standardized, sex-adjusted measurements. Classification functions derived from a stepwise multiple discriminant analysis correctly identified more than 87 percent of the specimens as being from either a northern or southern locality. These results support recognition of southern spotted dolphins as a separate management stock. Dolphins from a sample representing a far-western locality show similarities with those from the south. Patterns of geographic variation in morphology were evaluated using Mantel tests and matrix correlations; 17 of the 36 characters showed regional patterning and 18 exhibited local patterning. Morphological trends were also assessed with respect to 14 oceanographic measures that characterize environmental variability in the ETP. In general, clinal variation in morphology mirrors a north-south trend found in several environmental variables. Overall, the most striking associations between morphological and environmental variables involved Solar Insolation (Jan.). Sea surface temperatures also exhibited a strong association with morphological variables; the highest correspondence was found between width of the temporal fossa and sea surface temperature in July. In addition, the thickness of the oxygen minimum layer exhibited a pattern of variation that was statistically associated with 58 percent of the skull characters. The geographic patterns found for cranial morphology and environmental measures are strongly clinal and in general congruent, suggesting that oceanographic conditions in part determine trends in morphological characteristics of spotted dolphins.     

DEVELOPMENTAL PATTERN OF THE BONY FALX AND BONY TENTORIUM OF SPOTTED DOLPHINS (STENELLA ATTENUATA) AND THE RELATIONSHIP BETWEEN DEGREE OF DEVELOPMENT AND AGE

The bony falx (BF) and bony tentorium (BT) of spotted dolphins ( Stenella attenuata ) at different growth stages were observed. The BF and BT were observed to have the shape of the entire falx cerebri (FC) and tentorium cerebelli (TC), respectively, in adults, but were not seen in fetuses at all. Partially formed BF and BT were observed in the young. The BF and BT of spotted dolphins are formed by ossification in the FC and TC in the course of aging. Based on age determination by means of dentinal growth layers of the maxillary teeth, the ossification in the FC and TC of spotted dolphins begins by about 1 yr of age, and is completed, except for the passage of nerves and blood vessels, by about 9 yr of age. The BF and BT of spotted dolphins are formed possibly by continual dietary intake of large amounts of calcium, phosphorus and vitamin D.     

SOCIAL ECOLOGY OF BOTTLENOSE DOLPHINS IN THE KVARNERIĆ (NORTHERN ADRIATIC SEA)

A bottlenose dolphin community was studied from small inflatable craft from 1987 to 1994 in a relatively large area (about 800 km²) east of the islands of Loˇsinj and Cres, northern Adriatic Sea. A total of 106 individuals were photoidentified based on natural permanent marks on their dorsal fins. Most of the dolphins were resighted on a regular basis, indicating a high level of year-round site fidelity, although their range was evidently greater than the chosen study area. Dolphin density was highly variable and considerably lower than for most well-known bottlenose dolphin communities. Groups averaged seven individuals, with a mode of two. Groups entirely composed of adults were the smallest, groups with calves the largest. Group fluidity was high, seasonal and yearly changes in mean group size being also considerable. Summer was the peak calving season, with a striking variation in the number of births on alternate years. Poor evidence of shark predation was found. The social organization of this dolphin community seemed to be highly flexible, possibly as an adaptation to cope with environmental changes as well as with a limited and variable availability of prey.     

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.     

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.     

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.     

FOOD AND FEEDING ECOLOGY OF THE COMMON DOLPHIN (DELPHINUS DELPHIS) IN THE OCEANIC NORTHEAST ATLANTIC AND COMPARISON WITH ITS DIET IN NERITIC AREAS

The short-beaked common dolphin, Delphinus delphis , is abundant in both neritic and oceanic habitats. These two domains differ largely in terms of the nature of forage organisms and their availability to surface-bound top predators, which suggests that the common dolphin should show extensive variability in foraging strategies as a response to these different habitats. However, although its diet is well known over continental shelves, so far, mostly because of sampling issues, nothing has been published on its diet in oceanic habitats. In this study, the diet of sixty-three common dolphins bycaught in the French albacore tuna driftnet fishery in the summers of 1992–1993 in the oceanic Bay of Biscay was determined from their stomach contents and compared to neritic studies. The diet was dominated by fish (90% by number [N] and 53% by mass of total diet [M]). Cephalopods were also important in the total diet (9%N, 46%M) but were a prey of secondary importance in the fresh fraction (3%N, 10%M), presumably due to longer retention of cephalopod remains in the stomach. Crustaceans were of minor importance. At the species level, the myctophid fish ( Notoscopelus kroeyeri ) largely dominated the diet. Prey size ranged from 1 to 68 cm, but the majority of preys were from 2 to 30 cm. The prey characteristics and their state of digestion suggest that the common dolphin forages preferentially on small schooling, vertically migrating mesopelagic fauna in the surface layer at dusk and early night. The diet is taxonomically distinct from results obtained in neritic studies but is similar in terms of prey type and the corresponding feeding behavior.     

DIURNAL BEHAVIOR OF FREE-RANGING BOTTLENOSE DOLPHINS IN THE KVARNERIĆ (NORTHERN ADRIATIC SEA)1

The diurnal behavior of a bottlenose dolphin (Tursiops truncatus) community was observed from small inflatable craft between 1987 and 1994. Following a preliminary ad libitum study 11,839 3-min behavioral samples were recorded in 1991-1994. The behavioral budget showed a predominance (about 80%) of activities characterized by long (>30 sec) dives, considered to be largely related to prey search or feeding. Obvious foraging near the surface was observed rarely. The frequent following of trawlers (accounting for 4.6% of the behavioral budget) was indicative of the presence of alternative strategies for finding food. Yearly and seasonal behavioral variation-particularly in feeding-related and travel behaviors-was consistent with the hypothesis of behavioral flexibility as a response to environmental changes and fluctuating prey kind and availability. Yearly shifts in social behavior appeared to be partly influenced by breeding cycles. Groups engaged in feeding-related activities were significantly smaller than traveling or socializing groups, and dramatic interannual group-size shifts seemed to be largely affected by environmental variables, rather than being entirely determined by behavioral activity changes. The remarkable behavioral flexibility of this bottlenose dolphin community may contribute to its survival in the shifting environmental conditions of the northern Adriatic Sea. However, the high proportion of time consistently devoted to feeding-related activities, as compared to other areas, suggests that food resources in the Kvarneri? were not only highly variable but also depleted.