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.     

A VIDEO SONAR AS A NEW TOOL TO STUDY MARINE MAMMALS IN THE WILD: MEASUREMENTS OF DOLPHIN SWIMMING SPEED

The use of a multibeam sonar for fine-scale studies of the foraging activity of bottlenose dolphins ( Tursiops truncatus ) was tested in a coastal environment where a group of 14 individuals is known to be resident. The sonar was set on a rubber boat and the signals were received continuously as the dolphins were foraging in a pass swept by strong tidal currents. These signals were recorded in HI 8 video standard. The system produced real-time undistorted images of the dolphins and of their movements within the limits of the sonar beam (15° vertical width, 90° horizontal width, range set at 50 m). The average swimming speed relative to the bottom was 1.8 m ·sec⁻¹ but swimming speed relative to the water was 2.2 m-sec⁻¹ when allowance was made for the average 1.2 m·sec⁻¹ tidal current. Maximum swimming speed relative to the water was as high as 4.8 m·sec⁻¹. It is estimated from studies of swimming energetics that continuously foraging in a current vein flowing at 1.2 m·sec¹ would represent only small additional costs compared to foraging out of these currents. However, previous observations of the same pod have shown that these dolphins keep foraging in this site when current speed reaches 2.7 m·sec⁻¹; under such circumstances, the cost of foraging in this site was calculated to increase by 96%. This suggests that foraging efficiency should be dramatically enhanced during rising tide since the dolphins consistently forage in these currents year round.     

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.     

STABILIZATION MECHANISM IN SWIMMING ODONTOCETE CETACEANS BY PHASED MOVEMENTS

Propulsive movements of the caudal oscillating flukes produce large forces that could induce equally large recoil forces at the cranial end of the animal, and, thus, affect stability. To examine these vertical oscillations, video analysis was used to measure the motions of the rostrum, pectoral flipper, caudal peduncle, and fluke tip for seven odontocete cetaceans: Delphinapterus leucas, Globicephala melaena, Lagenorhynchus obliquidens, Orcinus orca, Pseudorca crassidens, Stenella plagiodon , and Tursiops truncatus. Animals swam over a range of speeds of 1.4–7.30 m/sec. For each species, oscillatory frequency of the fluke tip increased linearly with swimming speed. Peak-to-peak amplitude at each body position remained constant with respect to swimming speed for all species. Mean peak-to-peak amplitude ranged from 0.02 to 0.06 body length at the rostrum and from 0.17 to 0.25 body length at the fluke tip. The phase relationships between the various body components remain constant with respect to swimming speed. Oscillations of the rostrum were nearly in phase with the fluke tip with phase differences out of—9.4°-33.0° of a cycle period of 360°. Pectoral flipper oscillations trailed fluke oscillations by 60.9°-123.4°. The lower range in amplitude at the rostrum compared to the fluke tip reflects increased resistance to vertical oscillation at the cranial end, which enhances the animal's stability. This resistance is likely due to both active and passive increased body stiffness, resistance on the flippers, phased movements of body components, and use of a lift-based propulsion. Collectively, these mechanisms stabilize the body of cetaceans during active swimming, which can reduce locomotor energy expenditure and reduce excessive motions of the head affecting sensory capabilities.     

ORPHAN-INDUCED LACTATION IN TURSIOPS AND ANALYSIS OF COLLECTED MILK

Two non-lactating, non-gravid female bottlenose dolphins, Tursiops truncatus , (SLA, age 32 yr and TOD, age 34 yr) began lactating within one week after being housed with a 6.5- and 6.75-month-old orphaned calf, respectively. Cooperative swimming and nursing behaviors were quickly evident in both cow/ calf pairs. During the first 2.5 mo a milk substitute, eventually in combination with some whole fish, was used to supplement nursing in both calves. After this initial period, supplemental feeding was discontinued. SLA then became her adopted calf s sole source of nutrition for the next 5 mo, followed by another 6 mo during which the calf shifted to a predominantly fish diet with occasional nursing. TOD's orphan also continued to nurse beyond the 2.5-mo supplementation period and began voluntarily accepting small quantities of fish, suckling continued over the next year while fish consumption increased. Using a modified human breast pump, trainers collected milk at intervals from the adult dolphins throughout the lactation period. The first milk collected from TOD (day 8 after housing with calf) and SLA (day 12 with her calf) contained 6.0% and 10.3% fat, respectively. On day 68 TOD's milk fat had increased to 23.5%, and on day 37 that of SLA measured 22.5%. These later values are similar to those reported for normally lactating bottlenose dolphins.     

THE DIETS OF MODERN AND HISTORIC BOTTLENOSE DOLPHIN POPULATIONS REFLECTED THROUGH STABLE ISOTOPES

The δ¹³C and δ¹⁵N compositions of teeth used in combination with existing data provide dietary information for different populations of western North Atlantic bottlenose dolphins (Tursiops truncatus). The dental isotopic signatures of bottlenose dolphins collected during the 1980s significantly differ for coastal and offshore ecotypes and are consistent with reports that coastal forms feed primarily on fish whereas offshore individuals consume more squid. In a second study, the isotopic compositions of teeth from bottlenose dolphins that span a 100-yr period and data from published stomach content analyses as well as field observations made during the past 100 yr provide evidence that coastal bottlenose dolphins from the 1880s, 1920s, and 1980s had similar diets.     

THE DISTRIBUTION OF WHITE-SIDED (LAGENORHYNCHUS ACUTUS) AND COMMON DOLPHINS (DELPHINUS DELPHIS) VS. ENVIRONMENTAL FEATURES OF THE CONTINENTAL SHELF OF THE NORTHEASTERN UNITED STATES

Sightings of white-sided ( Lagenorhynchus acutus ) and common dolphins ( Delphinus delphis ) from survey vessels operating on the continental shelf of the northeastern United States were recorded during spring and fall of six years. Distributions of these sightings were compared with sea surface temperature and salinity, and bottom topography. The apparent geographic distributions of the two species are complementary, although there is a broad overlap. White-sided dolphins occurred primarily within the Gulf of Maine, but also on Georges Bank and in the mid-Atlantic region. Common dolphins were abundant within a broad band paralleling the continental slope (100–200 m depth contour) from latitude 35°N to the northeast peak of Georges Bank. Sightings of this species were distributed primarily along the edge of the continental shelf south of 40°N in spring and north of this latitude in fall. Few sightings of common dolphins occurred in the Gulf of Maine.
Although both species of dolphins were sighted more frequently in areas of high sea floor relief, white-sided dolphins occurred in areas where sea surface temperatures and salinities are low while common dolphins were sighted in warmer, more saline waters. However, these environmental conditions may be only secondarily influencing dolphin distribution. Seasonal variation in sea surface temperature and salinity, and local nutrient upwelling in areas of high sea floor relief may affect preferred prey abundances, which in turn may affect dolphin distribution.     

DOLPHIN THYROID AND ADRENAL HORMONES: CIRCULATING LEVELS IN WILD AND SEMIDOMESTICATED TURSIOPS TRUNCATUS, AND INFLUENCE OF SEX, AGE, AND SEASON

Biological and environmental influences on circulating adrenal and thyroid hormones were investigated in 36 wild and 36 semidomesticated Atlantic bottlenose dolphins, Tursiops truncatus, matched by age, sex, and time of year when the samples were collected. Serum concentrations of thyroxine (free [fT41 and total [tT4]), triiodothyronine (free [fT3], total [tT3], and total reverse [rT37), cortisol, and aldosterone were determined by radio-immunoassay, Wild female dolphins had significantly higher levels of tT4, fT4 and fT3, an effect that was possibly related to reproduction and lactation. Semidomesticated females had higher tT3 than their wild counterparts. fT4 declined with age in wild dolphins, whereas rT3 was greatest in the older animals. Cortisol and aldosterone were both higher in wild animals sampled after a variable interval of up to four hours after encirclement by capture net. The pattern of adrenal hormone release suggested a mild stress response. Levels of both adrenal hormones were low in semidomesticated dolphins conditioned to present voluntarily their tails for blood sampling, an approach that appears to yield specimens representative of resting values for these constituents.     

IMPROVING MANAGEMENT OF OVERLAPPING BOTTLENOSE DOLPHIN ECOTYPES THROUGH SPATIAL ANALYSIS AND GENETICS

In the Northwest Atlantic the distribution of coastal bottlenose dolphins ( Tursiops truncatus ) overlaps with that of the offshore ecotype. We hypothesized that the distribution of the two ecotypes could be delineated by depth and/or distance from shore, facilitating their identification during surveys. We obtained 304 skin biopsy samples and identified each as either coastal or offshore using analysis of mitochondrial DNA. We then interpreted the spatial distribution of coastal and offshore forms using spatial analysis. Using a Classification and Regression Tree (CART) analysis, we found a statistically significant break in ecotype distribution at 34 km from shore. In waters beyond 34 km from shore and deeper than 34 m, all bottlenose dolphins were of the offshore ecotype. Within 7.5 km of shore, all 65 samples were of the coastal ecotype. Between these two areas only nine samples were collected, so the genetic composition of bottlenose dolphins in this area remains poorly known. To enhance our understanding of the spatial distribution of the two ecotypes, future research should obtain more biopsy samples in this zone. Nevertheless, our results indicate that a conservative abundance estimate for the coastal ecotype could be generated from surveys of bottlenose dolphins within 7.5 km of shore.     

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.     

Factors influencing dolphin depredation in coastal fisheries of the northern Aegean Sea: Implications on defining mitigation measures

Dolphin interactions with coastal fisheries are of major concern, reportedly leading to gear damage, which increases the cost of coastal fishing globally and in the Mediterranean Sea. The aim of this study was to determine the effect that gear, target species, mesh size, depth, soaking duration, fishing area, and season have on net depredation frequency and to offer insights on possible mitigation solutions. From November 2013 to February 2016 we monitored 107 active coastal fishers in 22 ports of the northern Aegean Sea coastline, identified the main target species of the fishery and recorded the damages on gill nets and trammel nets caused by dolphins, mainly the common bottlenose dolphin (Tursiops truncatus). Quasi-binomial generalized linear models were used to determine the relationship between the examined factors and depredation frequency. The analysis revealed that the gears mostly depredated were gill nets and trammel nets with small mesh sizes, mainly targeting surmullet (Mullus surmuletus), red mullet (Mullus barbatus), common sole (Solea solea), European hake (Merluccius merluccius), and caramote prawn (Melicertus kerathurus). The probability of depredation was also significantly dependent on the fishing area.     

Two new species of symbiotic ciliates from the respiratory tract of cetaceans with establishment of the new genus Planilamina n. gen. (Dysteriida, Kyaroikeidae)

Examination of mucus discharged from the blowhole of Atlantic bottlenose dolphin (Tursiops truncatus) at Marine Life Oceanarium, Gulfport, Mississippi, and false killer whale (Pseudorca crassidens) and Atlantic bottlenose dolphin at SeaWorld Orlando, Orlando, Florida, using live observations and protargol impregnation revealed mixed infections of Kyaroikeus cetarius and two new species. Planilamina n. gen. is characterized by a C-shaped argentophilic band located along the laterally flattened margin of cell and extending from the cell apex to subposterior cone-shaped podite; a deep oral cavity containing one short preoral kinety, two circumoral kineties, seven to 13 infundibular kineties, and a cytostome; a broadly funnel-shaped cytopharynx reinforced by argentophilic fibers but without nematodesmata; closely packed postoral kinetofragments set in a pocket located anterior left of the podite; and somatic kineties as a right field closely situated at the right surface and a left field bordering the anterior left margin of the oral cavity. The type species for the genus, Planilamina ovata n. sp., is distinguished from its sister species Planilamina magna n. sp. by the following characteristics: body size (28-65 x 20-43 microm vs. 57-90 x 40-63 microm), number of right field kineties (38-55 vs. 79-99), and position of the anterior end of the leftmost kinety in the right somatic field (anterior one-third vs. mid-body). The morphogenesis of Planilamina ovata is similar to that of K. cetarius. The diagnosis of Kyaroikeidae is emended to accommodate the new genus.     

ISOTOPIC ANALYSIS OF δ13C, δ15N, AND δ34S "A FEEDING TALE" IN TEETH OF THE LONGBEAKED COMMON DOLPHIN, DELPHINUS CAPENSIS

Stable isotopic analyses of carbon, nitrogen, and sulfur were performed on teeth of different ages and sexes of the longbeaked common dolphin, Delphinus capensis, from the Gulf of California. Similarities in diet are suggested between the sexes, with no significant differences in isotopic compositions being observed. Differences in the δ¹³C, δ¹⁵N, and δ³⁴S signatures were found among the age groups (nursing calf, juvenile, subadult, and adult). These data suggest that this species is generally a coastal feeder, and that it changes its feeding habits with increasing age, drawing more nutrition from higher trophic level organisms later in life.     

东海水域瓶鼻海豚种群数量和分布的初步研究

运用截线抽样法调查了北纬２５°～３０°,东经１２５°以西的东海水域中瓶鼻海豚在夏季７～９月间的种群密度和数量,得出的最小估计值分别为０．１４００头／ｋｍ２和９９４０头。瓶鼻海豚在该水域的分布特点为：随纬度增加（由南向北）密度逐渐减少,而在经度方向上则没有明显的密度变化规律。     

MORBILLIVIRUS INFECTION IN BOTTLENOSE DOLPHINS: EVIDENCE FOR RECURRENT EPIZOOTICS IN THE WESTERN ATLANTIC AND GULF OF MEXICO

Morbillivirus infection is widespread among odontocetes of the western Atlantic and Gulf of Mexico. Serologic evidence of infection in bottlenose dolphins, Tursiops truncatus , was first detected during an epizootic along the mid-Atlantic coast in 1987. Here, we report recurrent epizootics in the coastal dolphin population since at least the early 1980s based on serological surveys and regional stranding frequencies. The first observed epizootic of this series occurred in the Indian and Banana Rivers in 1982 and was followed by others on the mid-Atlantic coast in 1987–1988 and in the Gulf of Mexico between 1992 and 1994. This temporal pattern of infection is likely facilitated by the population size and its fragmentation into relatively discrete coastal communities. Introduction of morbillivirus into a community with a sufficient number of naive hosts may precipitate an epizootic, depending on the potential for transmission within the group. Propagation of an epizootic along the coast is probably determined by frequency of contact between adjacent communities and seasonal migrations.
Morbillivirus antibodies were also detected in serum from offshore bottlenose dolphins. The sero-prevalence in the latter may be higher than in coastal dolphins because of their close association with enzootically infected pilot whales ( Globicephala spp.). Occasional contact between offshore and coastal dolphins may provide an epizootiologic link between pilot whales and coastal dolphin communities.     

LOCAL ABUNDANCE AND DISTRIBUTION OF BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) IN THE NEARSHORE WATERS OF VIRGINIA BEACH,VIRGINIA

We investigated patterns of abundance and distribution for coastal migratory Atlantic bottlenose dolphins (Tursiops truncatus) that appear seasonally in the nearshore waters of Virginia Beach, Virginia. The study was conducted along 24 km of shoreline at the southern point of the Chesapeake Bay mouth from April 1994 to March 1995. This is the first study to investigate the relationship between the abundance of coastal migratory dolphins and factors that might affect their movement. A profile analysis of variance revealed significant differences in local abundance and distribution throughout the year. Dolphin number was positively correlated with water temperature and not correlated with photoperiod. Although prey distribution and abundance are two factors thought to affect dolphin presence, in this study the relationship between these two factors and dolphin abundance was unclear. Greater numbers of dolphins were found in the ocean section of the study area. However, significantly higher ratios of neonatal dolphins were observed in the bay section, suggesting the bay serves as a nursery area. The observed relationship between local dolphin abundance and environmental factors in Virginia may provide insight into dolphin distribution and migration along the Atlantic coast of the United States.     

SHARK ATTACKS ON BOTTLENOSE DOLPHINS (TURSIOPS ADUNCUS) IN SHARK BAY, WESTERN AUSTRALIA: ATTACK RATE, BITE SCAR FREQUENCIES, AND ATTACK SEASONALITY

Shark predation may have been a central factor influencing the evolution of sociality in dolphins, as well as a determinant of dolphin habitat use and behavior. To understand the role of predation in driving interpopulation differences in behavior and sociality, it is important to quantify differences in predation risk among populations. This study describes the frequency of shark-inflicted scars and estimates the shark attack rate on bottlenose dolphins ( Tursiops aduncus ) in Shark Bay, Western Australia. Shark bite scars were found on 74.2% (95 of 128) of non-calves, and most of these scars were inflicted by tiger sharks ( Galeocerdo cuvier ). Although there were no differences among age/sex classes in the frequency of scarring, significantly more adult males than adult females bore multiple scars. The rate of unsuccessful shark attack was estimated to be between 11% and 13% of dolphins attacked each year. Large sharks (>3 m) were responsible for a disproportionate number of attacks. However, bites from small carcharhinid sharks on 6.2% of dolphins suggest that some of these small sharks may be dolphin ectoparasites. Both the scar frequencies and attack rate suggest that Shark Bay dolphins face a greater risk of predation than bottlenose dolphins in other locations.     

AGE STRUCTURE AND GROWTH OF THE BOTTLENOSE DOLPHIN (TURSIOPS TRUNCATUS) FROM STRANDINGS IN THE MISSISSIPPI SOUND REGION OF THE NORTH-CENTRAL GULF OF MEXICO FROM 1986 TO 2003

Despite their high abundance and wide distribution, little is known about the historical or current growth and age structure of coastal bottlenose dolphins ( Tursiops truncatus ) in the north-central Gulf of Mexico. Between 1986 and 2003, teeth were collected from bottlenose dolphins stranded on the mainland coast of Mississippi and the adjacent barrier islands. Bottlenose dolphin strandings occurred year round, peaking in March and April. Neonate strandings also peaked during these 2 mo. Age estimates were obtained from 111 animals by reading the growth layer groups in the dentine layer of the teeth. The ages ranged from <1 yr to 30 yr of age. The two-stage Laird–Gompertz growth model was fitted to the total length and age data. On the basis of this model, the asymptotic lengths were estimated at 250 cm for females and 255 cm for males. The length at birth estimates were 98–103 cm for females and 100–107 cm for males. These lengths are similar to those of bottlenose dolphin populations from other Gulf of Mexico areas and from the North Atlantic Ocean along the southeastern United States.     

LATERAL MANDIBULAR WALL THICKNESS IN TURSIOPS TRUNCATUS: VARIATION DUE TO SEX AND AGE

In odontocetes the mandibular bone serves two functions: to capture prey, and as a means of the reception and transmission of sound waves through a fat body in the mandibular canal, which opens posteriorly as the mandibular foramen. The posterior part of the lateral wall of the odontocete mandible is thin, and appears to represent a compromise between a strong mandible for prey capture and a thin vibrating plate for hearing. We studied the intraspecific variation of minimum thickness of the lateral mandibular wall along four transects (T1-T4) at the area of the mandibular foramen, in relation to the skull size and the mandibular size in different-aged bottlenose dolphins, Tursiops truncatus (18 females between 1 and 42 yr, 17 males between 1 and 32 yr). The minimum thickness was absolutely at its lowest at the most posterior transect T1, but did not vary significantly between the sexes or between the ages. The minimum thickness varied significantly at the two most anterior transects, T3 and T4, both between the sexes and among the ages. The thickness increased throughout life among males, whereas in females it first increased and then starts to decrease around the age of 20.     

BUFFERING CAPACITY OF THE LOCOMOTOR MUSCLE IN CETACEANS: CORRELATES WITH POSTPARTUM DEVELOPMENT, DIVE DURATION, AND SWIM PERFORMANCE

Skeletal muscles of marine mammals must support the metabolic demands of exercise during periods of reduced blood flow associated with the dive response. Enhanced muscle buffering could support anaerobic metabolic processes during apnea, yet this has not been fully investigated in cetaceans. To assess the importance of this adaptation in the diving and swimming performance of cetaceans, muscle buffering capacity due to non-bicarbonate buffers was measured in the longissimus dorsi of ten species of odontocete and one mysticete. Immature specimens from a subset of these species were studied to assess developmental trends. Fetal and neonatal cetaceans have low buffering capacities (range: 34.8–53.9 slykes) that are within the range measured for terrestrial mammals. A lengthy developmental period, independent of muscle myoglobin postnatal development, is required before adult levels are attained. Adult cetacean buffering capacities (range: 63.7–94.5 slykes) are among the highest values recorded for mammals. Cetacean species that demonstrate extremely long dive durations or high burst speed swimming tend to have greater buffering capacities. However, the wide range of body size across cetaceans may complicate these trends. Enhanced muscle buffering capacity may enable small-bodied species to extend breath-hold beyond short aerobic dive limits for foraging or predator evasion when necessary.