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An understanding of cetacean distribution is necessary to gain insights into crucial ecological processes for species conservation management. However, cetacean habitat preference and distribution in the northern Savu Sea, Indonesia are still poorly understood. We use maximum entropy modelling with five environmental predictors to describe habitat preference and distribution of seven cetacean species in the northern Savu Sea. Our study confirms that static predictors are the most important variables in explaining habitat preferences of seven cetacean species in the northern Savu Sea. Seasonally, each cetacean species has a different environmental preference. Globicephala macrorhynchus prefers the open nearshore areas adjacent to deep waters. Stenella attenuata is associated with nearshore habitats and higher productivity areas. Stenella longirostris prefers nearshore habitat during coastal upwelling events. Tursiops truncatus prefers nearshore sheltered habitat. Grampus griseus habitats are highly related to depth and steep gradients around the isobaths of 200-1,000 m. Feresa attenuata are associated with deep waters and cooler sea temperature. Pseudorca crassidens prefers protective bays with productive waters. This study provides important information for reviewing marine spatial plans of the northern Savu Sea, specifically on managing species fisheries interaction, optimizing surveillance, and regulating marine traffic.

Endemic subspecies and multiple stocks have been proposed for spinner (Stenella longirostris) and spotted dolphins (Stenella attenuata), two species with historically high mortality due to bycatch in the eastern tropical Pacific yellowfin tuna fishery. However, there has been low statistical power in tests for genetic differentiation among most of these stocks, due to large historical abundances, ongoing gene flow, and recent divergence. We tested for structure at multiple hierarchical levels by collecting whole mitochondrial genome sequences (mtDNA) and nuclear SNPs (nuDNA) from 104 spinner and 76 spotted dolphins using capture array library enrichment and highly paralleled DNA sequencing. MtDNA showed weak but significant differences between subspecies of spotted (FST: 0.0125; P = 0.0402) and spinner dolphins (FST: 0.0133; P = 0.034). NuDNA supported subspecies of spotted but not spinner dolphins. Relatively strong and significant differentiation was detected between whitebelly and eastern spinner stocks using nuDNA (FST: 0.0297; P = 0.0059). Neither mtDNA nor nuDNA supported the division of existing offshore stocks of spotted dolphins or Tres Marias spinner dolphins. This work identifies a genetic basis for biologically meaningful management units of these two species, a critical component in understanding their response in the face of historical and continued fishery interactions.

This project was funded by the Office of Naval Research (award number: N000141110612; Program Manager Michael J. Weise), WHOI Marine Mammal Center, and the Sawyer and Penzance Endowed Funds, with additional field time funded by grants through Cascadia Research Collective by the National Oceanographic Partnership Program (through the Alaska SeaLife Center) and the Pacific Islands Fisheries Science Center. PLT acknowledges the support of the MASTS pooling initiative (The Marine Alliance for Science and Technology for Scotland). MASTS is funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions. Postprint

Phylogenetic relationships in the family Delphinidae have been widely debated. We examined 347 skulls of Tursiops, Stenella, Delphinus, Steno, Lagenodelphis, and Sousa in order to resolve the phylogenetic position of Australian species of Tursiops. Five Tursiops type specimens were included. Cranial morphology was described using 2-dimensional (2-D) and 3-dimensional geometric morphometrics (3-GM), counts and categorical data. Analyses showed a clear morphological separation of Tursiops, including type specimens, from other genera. The three Stenella species did not cluster together. Stenella attenuata clustered with Delphinus delphis, and Stenella coeruleoalba with Lagenodelphis hosei. Length and width of the skull and rostrum were important discriminators in both methods. For 3-D data, round vs. angular posterior skull shape distinguished some genera. Taxa that overlapped in the multivariate analyses had different mean tooth counts. Our study challenges genetic studies that identified Tursiops as polyphyletic, with T. aduncus closer to S. attenuata.

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

Hematology, serum chemistry, and plasma hormones were evaluated in 72 pantropical spotted dolphins (Stenella attenuata attenuata) from the eastern tropical Pacific in an attempt to define the degree of stress associated with chase and encirclement by a tuna purse seiner, and are here reported for the first time for this species. Dolphins had high levels of dopamine and moderately elevated levels of enzymes indicative of the expected muscle damage following exertion of the chase. The length of time between the start of the capture operation and blood sampling correlated with increases in platelet and white blood cell counts and mean cell hemoglobin concentration, while the length of time between net tie-down and blood sampling influenced platelet, white blood cell, and eosinophil counts. Ten dolphins recaptured 1–3 d after their first capture had significantly lower serum creatinine kinase, thyroid (T4) and globulin levels compared to values in dolphins sampled at nominal first capture. Although small sample sizes and large individual variation limit interpretation, these data indicate a stress response occurred in all dolphins, but the extent of the response is within the expected range for adaptive responses previously measured in limited numbers of wild mammals.

Food is one of the most important dimensions of resource partitioning for species coexistence. In this study, we investigated the dietary composition and foraging habits of three sympatric odontocetes in order to identify their levels of food niche overlap and ecological separation. Stomach content analysis was performed on samples collected from carcasses confiscated by police or entangled in gill nets from 1994 to 2001, including 27 Risso's dolphins (GG) (Grampus griseus), 27 Fraser's dolphins (LH) (Lagenodelphis hosei), and 45 pantropical spotted dolphins (SA) (Stenella attenuata). GG consumed only cephalopods, with Enoploteuthis chunii accounting for 90.5% of total prey consumed, LH fed on mesopelagic fishes and cephalopods, dominated by hatchetfish, Polyipnus stereope (50.2%), and SA ate both mesopelagic and epipelagic preys, primarily fishes of Myctophum asperum (20.3%) and squids of E. chunii (25.8%). Among the three odontocetes, GG had the narrowest dietary niche width, while SA had the widest width. Both the niche overlap index and the analysis of similarities (ANOSIM) showed significant diet differentiation among these three dolphin species. The depth distribution of their principal prey items further suggests that LH feeds in the deepest waters while SA utilizes prey resources near surface.

Unlike other mammals, odontocetes and mysticetes have highly derived craniofacial bones. A growth process referred to as "telescoping" is partly responsible for this morphology. Here, we explore how changes in facial morphology during fetal growth relate to differences in telescoping between the adult odontocete Stenella attenuata and the mysticete Balaena mysticetus. We conclude that in both Stenella and Balaena head size increases allometrically. Similarly, odontocete nasal length and mysticete mouth size have strong positive allometry compared to total body length. However, the differences between odontocetes and mysticetes in telescoping are not directly associated with their fetal growth patterns. Our results suggest that cranial changes related to echolocation and feeding between odontocetes and mysticetes, respectively, begin during ontogeny before telescoping is initiated.

We radio tracked 20 pantropical spotted dolphins (Stenella attenuata )i n the pelagic waters of the eastern tropical Pacific Ocean during three research cruises in 1992, 1993, and 2001. Dolphins were tracked for up to 6 d from 20 to 1,000 km offshore. Seventeen of these dolphins also carried time–depth recorders, nine of which were recovered with a combined total of 477 h of dive data. The movement data suggested that the dolphins associate with areas of relatively high biological productivity. Dolphins close to shore moved along the continental slope, while some dolphins traveled farther offshore along thermocline “ridges.” Tracking and tagging multiple dolphins demonstrated the dynamic nature of dolphin herds, which change size and membership over the course of a day. At night, the dolphins traveled more slowly but dove deeper and longer, with more rapid ascents and descents than during daylight hours. These nighttime dive characteristics support the hypothesis suggested by some food habit studies that spotted dolphins are nocturnal feeders. Comparison of dive data with acoustic backscatter data indicates that spotted dolphins typically begin and end nighttime feeding activities with dusk and dawn diving bouts that track the vertical migration of organisms associated with the deep scattering layer.

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.

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.

The Gulf of Mexico is a subtropical marginal sea of the western North Atlantic Ocean with a diverse cetacean community. Ship-based, line-transect abundance surveys were conducted in oceanic waters (>200 m deep) of the northern Gulf within U.S. waters (380,432 km 2 ) during spring from 1996 to 1997 and from 1999 to 2001. Data from these five surveys were pooled and minimum abundance estimates were based on 12,162 km of effort and 512 sightings of at least 19 species. The most commonly sighted species (number of groups) were pantropical spotted dolphin, Stenella attenuata (164); sperm whale, Physeter macncephalus (67); dwarf/pygmy sperm whale, Kogia sitna/breviceps (58); Risso's dolphin, Grampus griseus (38); and bottlenose dolphin, Tursiops truncatus (24). The most abundant species (number of individuals; coefficient of variation) were S. attenuata (91,321; 0.16); Clymene dolphin, S. clymene (17,355; 0.65); spinner dolphin, S. longirostris (11,971; 0.71); and striped dolphin, S. coeruleoatha (6,505; 0.43). The only large whales sighted were P. macracephalus (1,349; 0.23) and Bryde's whale, Balaenoptera edeni (40; 0.61). Abundances for other species or genera ranged from 95 to 2,388 animals. Cetaceans were sighted throughout the oceanic northern Gulf and, whereas many species were widely distributed, some had more regional distributions.

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.

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.

We conducted a cetacean survey in the pelagic western tropical Indian Ocean (WTIO) aboard an 85-m research vessel from March to July 1995, covering 9,784 linear km. Using 25 × binoculars and line-transect methods, we recorded 589 sightings of 21 species. Stenella longirostris was the most abundant cetacean, in terms of number of individuals sighted, by an order of magnitude above any other species, while Physeter macrocephalus was the most frequently sighted, in terms of number of schools. Twelve species were widespread, seven were rare, and two were localized; our sightings include new distributional records for 12 species. Significant observations included the following: (1) Delphinus cf. tropicalis was abundant off the coast of Oman (16 sightings) and readily distinguishable in the field from D. delphis and D. capensis, (2) Balaenoptera musculus was fairly common and localized in the area of the Maldives (17 sightings), and (3) three sightings were made of an unidentified bottlenose whale tentatively referred to as Indopacetus (i. e., Mesoplodon) pacificus. We recorded 26 mixed-species cetacean schools, 43 schools with which seabirds associated, and 17 schools associated with tuna. Notable among these were mixed aggregations of Stenella attenuata, S. longirostris, yellowfin tuna, and seabirds. The cetacean community of the WTIO was similar to that of the eastern tropical Pacific (ETP) and the Gulf of Mexico (GM) in several respects. First, differences in abundance rank of individual species were small, with the result that common species were common and rare species were rare, regardless of ocean. Second, these differences in abundance were due primarily to differences in encounter rate, which varied with ocean by as much as 3,000%, and less so to school size, which generally varied less than 100%. Third, regardless of ocean, three species comprised the majority of cetaceans in the community, Stenella attenuata, S. longirostris, and S. coeruleoalba, representing 62%-82% of all individuals for all species. However, the rank order of abundance for these three species differed with ocean. Most notably, S. attenuata was abundant in the ETP and GM (abundance rank = 2 and 1, respectively) but much less common in the WTIO (abundance rank = 6). Although habitat preferences for S. attenuata appear to overlap considerably with those of S. longirostris in the ETP, our results suggest there may actually be significant differences between these two species. Detailed analysis of oceanographic correlates of distribution will be necessary in order to understand fully the habitat requirements of these pelagic dolphins, often the most conspicuous elements of tropical cetacean communities around the world.

The physical habitat of cetaceans found along the continental slope in the north-central and western Gulf of Mexico was characterized from shipboard sighting data, simultaneous hydrographic measurements, and satellite remote sensing. The study area was encompassed by the longitude of the Florida-Alabama border (87.5°W), the southernmost latitude of the Texas-Mexico border (26.0°N), and the 100-m and 2,000-m isobaths. Shipboard surveys were conducted seasonally for two years from April 1992 to May 1994. A total of 21,350 km of transect was visually sampled in an area of 154,621 km2. Sighting localities of species in the study area were differentiated most clearly with bottom depth. Atlantic spotted dolphins (Stenella frontalis) were consistently found in the shallowest water on the continental shelf and along the shelf break. In addition, the bottom depth gradient (sea floor slope) was less for Atlantic spotted dolphins than for any other species. Bottlenose dolphins (Tursiops truncatus) were found most commonly along the upper slope in water significantly deeper than that for Atlantic spotted dolphins. All the other species and species categories were found over deeper bottom depths; these were Risso's dolphins (Grampus griseus), short-finned pilot whales (Glob-icephala macrorhynchus), pygmy/dwarf sperm whales (Kogia spp.), roughtoothed dolphins (Steno bredanensis), spinner dolphins (Stenella longirostris), sperm whales (Physeter macrocephalus), striped dolphins (Stenella coeruleoalba), Mesoplodon spp., pantropical spotted dolphins (Stenella attenuata), Clymene dolphins (Stenella clymene) and unidentified beaked whales (Ziphiidae). Risso's dolphins and short-finned pilot whales occurred along the upper slope and, as a subgroup, were significantly different from striped dolphins, Mesoplodon spp., pantropical spotted dolphins, Clymene dolphins, and unidentified beaked whales, which occurred in the deepest water. Pygmy/dwarf sperm whales, rough-toothed dolphins, spinner dolphins, and sperm whales occurred at intermediate depths between these two subgroups and overlapped them.

Ontogeny and variation of head stripes were investigated in Stenella attenuata, S. longirostris, and Delphinus delphis. The eye stripe and the blowhole stripe together comprise the bridle. The Anlage of the bridle is a line extending across the back of the head in small fetuses. As the blowhole moves posteriorly and the rostrum lengthens, an inflection point develops in the line above the eye. This point moves forward, creating a progressively greater triangular area between the incipient eye and blowhole stripes. Well before birth, the inflection point is close to the apex of the melon, and the eye and blowhole stripes are narrow and well defined. This process proceeds to varying degrees in the other delphinoids. For example, in Peponocephala electra and in Cephalorhynchus spp., the inflection point migrates only part of the way to the apex of the melon, and the resulting “blowhole stripe” remains a broad caplike feature, resembling the state of development in small fetuses of Stenella and Delphinus. A simple eye spot without a stripe may be primitive, as it occurs in non-delphinoid cetaceans. Presence of the bridle only in the delphinids and phocoenids may unite them in a clade exclusive of the monodontids.

A method is presented for estimating age-specific mortality based on minimal information: a model life table and an estimate of longevity. This approach uses expected patterns of mammalian survivorship to define a general model of agespecific mortality rates. One such model life table is based on data for northern fur seals (Callorhinus ursinu.r) using Siler’s ( 1979) 5-parameter competing risk model. Alternative model life tables are based on historical data for human females and on a published model for Old World monkeys. Survival rates for a marine mammal species are then calculated by scaling these models by the longevity of that species. By using a realistic model (instead of assuming constant mortality), one can see more easily the real biological limits to population growth. The mortality estimation procedure is illustrated with examples of spotted dolphins (Stenella attenuata) and harbor porpoise (Phocoena phocoena).

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.

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. When studying morphological variation within a species, it can be important and informative to apportion variability into geographic and non-geographic components. Genetically controlled, non-geographic variation manifests itself in several ways, each of which may have distinct evolutionary implications. Sexual

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.