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.     

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.     

SEXUAL DIMORPHISM IN SPOTTED DOLPHINS (STENELLA ATTENUATA) IN THE EASTERN TROPICAL PACIFIC OCEAN

We analyzed 4 meristic and 32 morphometric cranial traits of 612 adult specimens of spotted dolphins ( Stenella attenuata ) from the eastern tropical Pacific Ocean for sexual dimorphism. Eleven 5" latitude/longitude blocks with five or more of each sex were assessed with a two-way ANOVA for sex and geographic differences. Interaction between these two factors was found for six measurements, suggesting that the degree of sexual dimorphism varies geographically for a few characters (although in no discernible geographic pattern). Sexual dimorphism was demonstrated for 23 of the 36 characters, with differences ranging from 0.00 to 5.88 percent. Females characteristically had a longer rostrum, while males generally had larger skulls overall. In terms of number of characters, the extent of sexual dimorphism demonstrated for skulls of spotted dolphins goes considerably beyond that shown for any other small delphine. A discriminant function involving a combination of 10 characters enabled us to identify correctly the sex of more than 75 percent of the specimens. A procedure for correcting specimen measurements is outlined that would enable an investigator to combine male and female specimens in geographic variation studies. A term (zwitter) is proposed for use when referring to specimens where measurements have been corrected to take into account differences between the sexes.     

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.     

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.     

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.     

ACOUSTIC IDENTIFICATION OF NINE DELPHINID SPECIES IN THE EASTERN TROPICAL PACIFIC OCEAN

Acoustic methods may improve the ability to identify cetacean species during shipboard surveys. Whistles were recorded from nine odontocete species in the eastern tropical Pacific to determine how reliably these vocalizations can be classified to species based on simple spectrographic measurements. Twelve variables were measured from each whistle ( n = 908). Parametric multivariate discriminant function analysis (DFA) correctly classified 41.1% of whistles to species. Non-parametric classification and regression tree (CART) analysis resulted in 51.4% correct classification. Striped dolphin whistles were most difficult to classify. Whistles of bottlenose dolphins, false killer whales, and pilot whales were most distinctive. Correct classification scores may be improved by adding prior probabilities that reflect species distribution to classification models, by measuring alternative whistle variables, using alternative classification techniques, and by localizing vocalizing dolphins when collecting data for classification models.     

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.     

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.     

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.     

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.     

GEOGRAPHIC VARIATION IN RATES OF VOCAL PRODUCTION OF FREE-RANGING BOTTLENOSE DOLPHINS

Echolocation and whistle production, group sizes, and activities of free-ranging bottlenose dolphins were compared across four regions (Wilmington, NC Intracoastal Waterway [ICW]; Wilmington coastline; Southport, NC coastline; and Sarasota, FL inshore waters). Number of whistles and echolo-cation bouts differed significantly across sites. Dolphins whistled significantly more in Southport than in the other sites, independent of group size. Unlike at the other sites, dolphin vocalizations in Southport did not vary significantly across activities; this difference may be due to the fact that Southport animals were often found behind shrimp-trawling vessels, which may affect their behavior. Resident Sarasota dolphins vocalized significantly less than dolphins at the NC sites. At most sites, echolocation production per dolphin decreased as group size increased, supporting the idea that echolocation information is shared. In the ICW and Sarasota, echolocation production per dolphin was highest while feeding, indicating that echolocation is used in foraging. At all sites but Southport, whistle production per dolphin was highest while socializing, indicating that whistles are used in communication. Overall, these data show that dolphins have different vocal and activity patterns at different sites; thus, caution should be used when extrapolating results from one study site to another.     

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.     

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.     

GEOGRAPHIC VARIATION IN SKULL MORPHOLOGY OF ADULT STELLER SEA LIONS (EUMETOPIAS JUBATUS)

Data from cranial specimens of adult E. jubatus were analyzed to compare intraspecific morphology of skulls. Males and females were grouped separately to avoid bias from sexual dimorphism. Geographic variation was observed in adult male E. jubutus , indicating the potential presence of three morphologically disparate groups: those from Alaska, those from California, and those from Japan and Russia. Although sample sizes were small, results from cluster and discriminant function analyses indicated that specimens from eastern and western Alaska were morphologically similar, and that the most divergent specimens for the species appeared to be those from Japan. Skulls from Alaska possessed a typically longer, less robust skull, whereas those from Japan appeared smaller, yet most robust. Skulls from California were intermediate.     

GEOGRAPHIC VARIATION IN THE LIFE HISTORY OF MASTOCARPUS PAPILLATUS (RHODOPHYTA)1

Variation in the geographic distribution of the life histories of Mastocarpus papillatus was investigated. Carpospores were isolated from 377 female gametophytes collected from eight localities on the Pacific coast of Baja California, Mexico, and California, U.S.A., and grown in laboratory culture. All carpospores from a single female gave rise either to basal discs with gametophyte-like uprights or crustose plants formerly referred to the genus Petrocelis. Early stages in the development of each type of germling were observed, and environmental factors affecting development were suggested. Based on carpospore germlings, females from each location were scored as having either the 1) sexual life history (crustose germlings) or 2) direct-development life history (discoid germlings with uprights). All females from the two southernmost locations in Baja California exhibited the sexual life history. In the three locations from the central-southern California coast, 70-95% of the females exhibited the sexual life history and the remainder exhibited the direct-development life history. In two of the three populations from the central-northern California coast, 70-90% of the females exhibited the direct-development life history and the remainder the sexual life history. In the third location from the central-northern California coast, the northernmost location sampled in the current study, 60% of the females exhibited the sexual life history and 40% the direct-development life history. The relative ecological advantages and disadvantages of the life histories are unknown as are the environmental factors that produced the ratios of sexual to direct-development females observed at each location.     

BLUE WHALE (BALAENOPTERA MUSCULUS)DISTRIBUTION IN THE EASTERN TROPICAL PACIFIC

The distribution of blue whales, Balaenoptera musculus , in the eastern tropical Pacific (ETP) was analyzed from 211 sightings of 355 whales recorded during research vessel sighting surveys or by biologists aboard fishing vessels. Over 90% of the sightings were made in just two areas: along Baja California, and in the vicinity of the Costa Rica Dome (a large, stationary eddy centered near 9°N, 89°W), with the rest made along the equator near the Galapagos islands, the coasts of Ecuador and northern Peru. All sightings occurred in relatively cool, upwelling-modified waters. Because these areas are the most productive parts of the ETP, and have relatively large standing stocks of euphausiids, it seems possible that blue whales select low latitude habitats which permit foraging. The waters off western Baja California were occupied seasonally, with a peak in sightings coinciding with the spring peak in upwelling and biological production. The Costa Rica Dome area was occupied year round, suggesting either a resident population, or that both northern and southern hemisphere whales visit, with temporal overlap. The modal group size was one for all areas and seasons, but the frequency of groups with two or more whales was significantly higher in sightings made near the Galapagos Islands and the coast of Ecuador and northern Peru.     

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.     

GEOGRAPHIC VARIATION AND SEXUAL DIMORPHISM IN THE SKULL OF DALL'S PORPOISE, PHOCOENOIDES DALLI

Abstract: Two hundred and eighty-nine skulls of Dall's porpoises from the North Pacific Ocean, Sea of Japan, Sea of Okhotsk, and Bering Sea were investigated morphometrically. A clear pattern of geographic variation was found in overall size of the skull. In the North Pacific, overall skull size became gradually smaller eastward from the coast of Japan to the offshore eastern Pacific, and it became larger again off the coast of California. Specimens from the Sea of Japan-Okhotsk and Bering Sea had larger skulls than those from the central North Pacific. The distribution of primary productivity corresponds well with this pattern of geographic variation, suggesting that quantity of food might affect the overall size of Dall's porpoise skulls. By canonical discriminant analyses, about 70% (male) and 90% (female) of specimens of the Sea of Japan-Okhotsk population were distinguished from those of other populations. This agrees with the results of genetic studies.