DIVE CHARACTERISTICS OF SATELLITE-MONITORED BLUE WHALES (BALAENOPTERA MUSCULUS) OFF THE CENTRAL CALIFORNIA COAST

Dive habits of four Northeast Pacific blue whales ( Balaenoptera musculus ) were studied using satellite-monitored radio tags. Tags summarized dive-duration data into eight 3-h periods daily. One tag additionally summarized dive depth and time-at-depth information for these same periods. Tracking periods ranged from 0.6 to 12.7 d and provided data for 17 three-hour summary periods, representing 2,007 dives (788 of which provided depth information). Total number of dives during a 3-h summary period ranged from 83 to 128. Seventy-two percent of dives were ≤ 1 min long. All whales spent >94% of their time submerged. Average duration of true dives (dives >1 min) ranged from 4.2 to 7.2 min. Seventy-five percent of depth-monitored dives were to ≤16 m, accounting for 78% of that animal's time. Average depth of dives to >16 m was 105 ± 13 m.     

AN ACOUSTIC LINK BETWEEN BLUE WHALES IN THE EASTERN TROPICAL PACIFIC AND THE NORTHEAST PACIFIC1

Blue whale calls in the eastern North Pacific Ocean consist of a two-part call often termed the A-B call. This call has been described for regions offshore of Oregon, Washington, and California, USA and the Sea of Cortez, Mexico (reviewed in Rivers 1997). Data collected from moored hydrophones in the eastern tropical Pacific (ETP) indicate that the A-B pattern is common in this region as well. There is consistency in this call type throughout the eastern North Pacific and throughout the year. This acoustic evidence indicates continuity between blue whales in the ETP and those found west of North America. The acoustic data suggest that the population of blue whales generally referred to as the “Californi/Mexico” stock might better be termed the “northeast Pacific” stock of blue whales.     

HISTORICAL OBSERVATIONS OF HUMPBACK AND BLUE WHALES IN THE NORTH ATLANTIC OCEAN: CLUES TO MIGRATORY ROUTES AND POSSIBLY ADDITIONAL FEEDING GROUNDS

The seasonal distributions of humpback and blue whales ( Megaptera novaeangliae and Balaenoptera musculus , respectively) in the North Atlantic Ocean are not fully understood. Although humpbacks have been studied intensively in nearshore or coastal feeding and breeding areas, their migratory movements between these areas have been largely inferred. Blue whales have only been studied intensively along the north shore of the Gulf of St. Lawrence, and their seasonal occurrence and movements elsewhere in the North Atlantic are poorly known. We investigated the historical seasonal distributions of these two species using sighting and catch data extracted from American 18th and 19th century whaling logbooks. These data suggest that humpback whales migrated seasonally from low-latitude calving/ breeding grounds over a protracted period, and that some of them traveled far offshore rather than following coastal routes. Also, at least some humpbacks apparently fed early in the summer west of the Mid-Atlantic Ridge, well south of their known present-day feeding grounds. In assessing the present status of the North Atlantic humpback population, it will be important to determine whether such offshore feeding does in fact occur. Blue whales were present across the southern half of the North Atlantic during the autumn and winter months, and farther north in spring and summer, but we had too few data points to support inferences about these whales' migratory timing and routes.     

INTERCHANGE AND ISOLATION OF HUMPBACK WHALES OFF CALIFORNIA AND OTHER NORTH PACIFIC FEEDING GROUNDS

Humpback whales feed in several high-latitude areas of the North Pacific. We examined the interchange of humpback whales between one of these areas, off California, and those in other feeding grounds in the eastern North Pacific:. Fluke photographs of 597 humpback whales identified off California between 1986 and 1992 were compared with those off Oregon and Washington (29); British Columbia (81); southeastern Alaska (343); Prince William Sound, Alaska (141); Kodiak Island, Alaska (104); Shumagin Islands, Alaska (22); and in the Bering Sea (7). A high degree of interchange, both inter-and intrayear, was found among humpback whales seen off California, Oregon, and Washington., A low rate of interchange was found between British Columbia and California.: two whales seen near the British Columbia/Washington border were photographed off California in a different year, No interchange was found between California and the three feeding areas in Alaska. Humpback whales off California, Oregon, and Washington form a single intermixing feeding aggregation with only limited interchange with areas farther north. These findings are consistent with photographic identification studies in the North Atlantic and with genetic studies in both the North Atlantic and North Pacific.     

BLUE WHALE VISUAL AND ACOUSTIC ENCOUNTER RATES IN THE SOUTHERN CALIFORNIA BIGHT

The relationship between blue whale ( Balaenoptera musculus ) visual and acoustic encounter rates was quantitatively evaluated using hourly counts of detected whales during shipboard surveys off southern California. Encounter rates were estimated using temporal, geographic, and weather variables within a generalized additive model framework. Visual encounters (2.06 animals/h, CV = 0.10) varied with subregion, Julian day, time of day, and year. Acoustic encounters of whales producing pulsed A and tonal B call sequences (song; 0.65 animals/h, CV = 0.06) varied by Julian day, survey mode (transit or stationary), and subregion, and encounters of whales producing downswept (D) calls (0.41 animals/h, CV = 0.09) varied by Julian day and the number of animals seen. Inclusion of Julian day in all models reflects the seasonal occurrence of blue whales off southern California; however, the seasonal peak in visual encounters and acoustic encounters of D calling whales (July–August) was offset from the peak in acoustic encounters of singing whales (August–September). The relationship between visual and acoustic encounter rates varied regionally, with significant differences in several northern regions. The number of whales heard D calling was positively related to the number of animals seen, whereas the number of singing whales was not related to visual encounter rate.     

ABUNDANCE OF BLUE AND HUMPBACK WHALES IN THE EASTERN NORTH PACIFIC ESTIMATED BY CAPTURE-RECAPTURE AND LINE-TRANSECT METHODS

We estimated humpback and blue whale abundance from 1991 to 1997 off the west coast of the U. S. and Mexico comparing capture-recapture models based on photographically identified animals and line-transect methods from ship-based surveys. During photo-identification research we obtained 4,212 identifications of 824 humpback whales and 2,403 identifications of 908 blue whales primarily through non-systematic small-boat surveys along the coast of California, Oregon, and Washington. Line-transect surveys from NOAA ships in 1991, 1993, and 1996 covered approximately 39,000 km along the coast of Baja California, California, Oregon, and Washington out to 555 km from shore. The nearshore and clumped distribution of humpback whales allowed photographic identification from small boats to cost-effectively sample a substantial portion of the population, but made it difficult to obtain effective samples in the line-transect surveys covering broad areas. The humpback capture-recapture estimates indicated humpback whale abundance increased over the six years (from 569 to 837). The broader more offshore distribution of blue whales made it harder to obtain a representative sample of identification photographs, but was well suited to the line-transect estimates. The line-transect estimates, after correction for missed animals, indicated approximately 3,000 blue whales (CV = 0.14). Capture-recapture estimates of blue whales were lower than this: approximately 2,000 when using photographs obtained from the line-transect surveys as one of the samples. Comparison of the results from the two methods provides validation, as well as insight into potential biases associated with each method.     

BLUE WHALE, BALAENOPTERA MUSCULUS, VOCALIZATIONS FROM THE WATERS OFF CENTRAL CALIFORNIA

Low-frequency calls produced by blue whales, Balaenoptera musculus , were recorded in the northeastern Pacific Ocean off central California. Two blue whales were sighted during a vessel-based marine mammal survey, and when sonobuoys were subsequently deployed, blue whale calls were recorded. A third recording was obtained during the survey from a blue whale that was not seen. Recordings with 15, 25, and 55 min of calls were obtained from these individuals. The three recordings all contain two-part, low-frequency calls with slight interindividual variation. The calls consist of an amplitude modulated (AM) signal with a mean center frequency of 16.5 Hz, followed by a downsweep whose mean center frequency sweeps from 18.2 Hz to 16.6 Hz. The recordings are compared with blue whale recordings from the Pacific and Atlantic Oceans. The geographic variability suggests that blue whale calls may be used as an acoustic indicator of stock identity.     

MOVEMENTS AND POPULATION STRUCTURE OF HUMPBACK WHALES IN THE NORTH PACIFIC

Despite the extensive use of photographic identification methods to investigate humpback whales in the North Pacific, few quantitative analyses have been conducted. We report on a comprehensive analysis of interchange in the North Pacific among three wintering regions (Mexico, Hawaii, and Japan) each with two to three subareas, and feeding areas that extended from southern California to the Aleutian Islands. Of the 6,413 identification photographs of humpback whales obtained by 16 independent research groups between 1990 and 1993 and examined for this study, 3,650 photographs were determined to be of suitable quality. A total of 1,241 matches was found by two independent matching teams, identifying 2,712 unique whales in the sample (seen one to five times). Site fidelity was greatest at feeding areas where there was a high rate of resightings in the same area in different years and a low rate of interchange among different areas. Migrations between winter regions and feeding areas did not follow a simple pattern, although highest match rates were found for whales that moved between Hawaii and southeastern Alaska, and between mainland and Baja Mexico and California. Interchange among subareas of the three primary wintering regions was extensive for Hawaii, variable (depending on subareas) for Mexico, and low for Japan and reflected the relative distances among subareas. Interchange among these primary wintering regions was rare. This study provides the first quantitative assessment of the migratory structure of humpback whales in the entire North Pacific basin.     

MIGRATION AND POPULATION STRUCTURE OF NORTHEASTERN PACIFIC WHALES OFF COASTAL BRITISH COLUMBIA: AN ANALYSIS OF COMMERCIAL WHALING RECORDS FROM 1908-1967

Data recorded from 24,862 whales killed by British Columbia coastal whaling stations between 1908 and 1967 revealed trends in the abundance, sex ratios, age structure, and distribution of sperm ( Physeter macrocephalus ), fin ( Balaenoptera physalus ), sei ( Balaenoptera borealis ), humpback ( Megaptera novaeangliae ), and blue ( Balaenoptera musculus ) whales. The catch data were analyzed using annual and monthly mean values. Monthly and annual variation in whaling effort was deduced from accounts of the history of British Columbia coastal whaling, and biases arising from changes in effort were considered in the interpretation of the results. During the later years of whaling (1948-1967), the mean lengths of captured whales declined significantly and pregnancy rates dropped to near zero in fin, sei, and blue whales. Monthly patterns in numbers killed revealed a summer migration of sei and blue whales past Vancouver Island, and confirms anecdotal suggestions that local populations of fin and humpback whales once spent extended periods in the coastal waters of British Columbia. Furthermore, the data strongly suggest that sperm whales mated (April-May) and calved (July-August) in British Columbia's offshore waters. The historic whaling records reveal much about the migratory behavior and distribution of the large whales species as they once were, and may continue to be, in the northeastern Pacific. Verifying the persistence of these trends in the remnant populations is a necessary and logical next step.     

FUNCTIONAL REDUCTION OF THE SOUTHERN MINKE WHALE (BALAENOPTERA ACUTOROSTRATA) TESTIS DURING THE FEEDING SEASON

Seasonal variation in reproductive activity in the male southern minke whales was investigated. The animals were killed and collected during the feeding season (December-March) in the Antarctic Ocean in the eastern part of Area III (35°E-70°E, south of 60°S) and Area IV (70°E-130°E, south of 60°S). Blood samples, testes, epididymides, and vasa deferentia were collected from 62 males, which had testes weighing over 400 g each. Changes in testicular morphology and plasma testosterone, estradiol-17β and LH concentrations measured by enzyme immunoassay were investigated. Reduction of testicular function associated with different capture periods was found in the summer season. From December to March, body length and body weight remained steady, but decreases in testicular weight, epididymal weight, and testicular volume were found in February. During the same period, plasma testosterone concentration also declined. A significant decrease in the number of germ cells continued during the period of feeding season. An increase in area of seminiferous tubule tended to proceed in a number of germ cells. These changes reflected the percentage of spermatozoa in the vasa deferentia ; in particular, motile spermatozoa were observed in December. Based on morphological observation, spermatogenesis had also declined. These results indicate regulation of testicular function by testosterone, as in terrestrial mammals. A gradual decrease of testicular function occurred during December-January.     

A NEW HYBRID BETWEEN A BLUE WHALE, BALAENOPTERA MUSCULUS, AND A FIN WHALE, B. PHYSALUS: FREQUENCY AND IMPLICATIONS OF HYBRIDIZATION

A female hybrid between a fin ( Balaenoptera physalus ) and a blue whale ( B. mnusculus ) was caught in whaling operations in 1984 off northwestern Spain. Its coloration and body proportions were intermediate between those of a fin and a blue whale, although it was anomalously large (19.4 m) when compared to fin whales of similar age (4 yr). It was sexually immature, concomitant with its age but not its length if it were a fin whale. Molecular analyses revealed that the mother of the hybrid was a blue whale and the father a fin whale. Examination of data for the five fin-blue whale hybrids in the literature, plus other anecdotal reports, indicates that hybridization between these two species occurs in various geographic regions and is relatively frequent, notably in light of the absence of reported hybrids between other mysticetes. Either species may act as father or mother, and there does not appear to be a selection for a given sex among the hybrids. The reproductive capacity of these hybrids remains unknown, although incidence of reproductive impairment appears to be higher in hybrid males than in hybrid females.     

TWO TYPES OF BLUE WHALE CALLS RECORDED IN THE GULF OF ALASKA

At one time blue whales were found throughout the Gulf of Alaska, however, none have been sighted there in post-whaling era surveys. To determine if blue whales ( Balaenoptera musculus ) might now occur in the Gulf of Alaska, an array of hydrophones was deployed there in October 1999. Data were retrieved in May 2000 and in June 2001. Spectrograms from a random subsample comprising 15% of the ∼63,000 h of data were visually examined for blue whale calls. Call types attributed to both northeastern and northwestern Pacific blue whales were recorded. Both of these call types were recorded seasonally from the initial deployment date in October 1999 through the third week of December 1999 and then from July 2000 through mid-December 2000. Both call types were regularly recorded on the same hydrophone at the same time indicating clear temporal and spatial overlap of the animals producing these calls. Two blue whale call types were recorded in the Gulf of Alaska suggesting that perhaps two stocks use this area. The northeastern call type has now been documented from the equator up to at least 55°N in the eastern North Pacific.     

SWIMMING SPEEDS OF SINGING AND NON-SINGING HUMPBACK WHALES DURING MIGRATION

Limited data exist on swimming speeds of humpback whales ( Megaptera novaeangliae ) and none on swimming speeds of singing whales during migration. We tracked humpback whales visually and acoustically during migration from the breeding grounds past our study site on the east coast of Australia (latitude 26°28'S). The mean swimming speed for whales while singing was 2.5 km/h, significantly less than for non-singing whales with a mean of 4.0 km/h but significantly greater than the mean of 1.6 km/h observed for singing whales on the Hawaiian breeding grounds. Between song sessions, there was no significant difference in speeds between whales that had been singing and other whales. Migration speeds were less for whales while singing but increased during the season. Although humpback whales can swim rapidly while singing (maximum observed 15.6 km/h), they generally do not do so, even during migration. Slower migration by singers would delay their return to the polar feeding areas and may be costly, but may be a strategy to provide access to more females.     

SEPARATING SOUTHERN BLUE WHALE SUBSPECIES BASED ON LENGTH FREQUENCIES OF SEXUALLY MATURE FEMALES

When sexually mature, Antarctic (true) blue whales are substantially longer than pygmy blue whales. To estimate the proportions of these two subspecies in various regions, Bayesian mixture models were fitted to catch length frequencies of sexually mature females. The extent of rounding to 5-ft intervals was also estimated. Antarctic blue whales dominated (99.2%) pelagic catches south of 52°S, whereas pygmy blue whales dominated (99.9%) north of 52°S and in 35°–180°E. South of 60°S, only 0.7% (95% credibility interval 0.5%–1.0%) were pygmy blue whales, lower than the 7% upper bound currently assumed. Shore-based catches from SW Africa and those before 1937 from South Georgia and the South Shetlands were estimated to contain 90%–92% Antarctic blue whales. Actual proportions were probably higher, but these data show evidence of rounding (up to 19% of records), poor length-estimation methods, and other problems. The mean length of sexually mature female Chilean blue whales (77.1 ft, 23.5 m) was intermediate between pygmy (68.9 ft, 21.0 m) and Antarctic blue whales (83.4–86.3 ft, 25.4–26.6 m). A good fit to these data was obtained only by assuming that the Chilean whales are a separate subspecies or distinctive population. This finding is also consistent with their discrete distribution, and genetic and call type differences, compared to Antarctic and pygmy blue whales.     

PHYLOGENETIC RELATIONSHIPS OF BRYDE'S WHALES IN THE WESTERN NORTH PACIFIC AND ADJACENT WATERS INFERRED FROM MITOCHONDRIAL DNA SEQUENCES1

To clarify phylogenetic relationships of Bryde's whales, we examined the nucleotide sequence of the mitochondrial control region and cytochrome b gene in 33 animals: 12 from offshore waters of the western North Pacific, five from off the Solomon Islands, and 16 from the East China Sea and coastal waters of Kochi in southwestern Japan. For reference purposes, homologous sequences from four Balaenoptera species including four Bryde's whales collected in the eastern Indian Ocean were added. We found whales from the three sampling areas to be genetically distinct. The control region sequences suggested that the whales from the three areas separate at higher than the populational level from one another. The cytochrome b data indicated that genetic differences between whales off the Solomon Islands and animals in the other two areas are equivalent to values found among recognized Balaenoptera species, although such a relationship was not observed between the other two areas. We conclude that whales in the East China Sea and coastal waters of Kochi separate from Bryde's whales in offshore waters of the western North Pacific at higher than the populational level but lower than the specific level (i. e., at the subspecific level) and that whales off the Solomon Islands do not belong genetically to the Bryde's whale as previously recognized.     

DIETS OF BAIRD'S BEAKED WHALES, BERARDIUS BAIRDII, IN THE SOUTHERN SEA OF OKHOTSK AND OFF THE PACIFIC COAST OF HONSHU, JAPAN

Stomach contents were analyzed from 127 Baird's beaked whales, Berardius bairdii , taken in coastal waters of Japan. During late July-August of 1985–1987, 1989, and 1991, 107 samples were collected from off the Pacific coast of Honshu. An additional 20 samples were collected from whales taken in the southern Sea of Okhotsk during late August—September of 1988 and 1989. Prey identification using fish otoliths and cephalopod beaks revealed the whales fed primarily on deep-water gadiform fishes and cephalopods in both regions. Prey species diversity and the percentage of cephalopods and fish differed between the two regions. Off the Pacific coast of Honshu the whales fed primarily on benthopelagic fishes (81.8%) and only 18.0% on cephalopods. Eight species of fish representing two families, the codlings (Moridae) and the grenadiers (Macrouridae), collectively made up 81.3% of rhe rotal. Thirty species of cephalopods representing 14 families made up 12.7%. In the southern Sea of Okhotsk, cephalopods accounted for 87.1% of stomach contents. The families Gonatidae and Cranchiidae were the predominant cephalopod prey, accounting for 86.7% of the diet. Gadiform fish accounted for only 12.9% of the diet. Longfin codling, Laemonema longipes , was the dominant fish prey in both regions. Depth distribution of the two commonly consumed fish off the Pacific coast of Honshu indicate the whales in this region fed primarily at depths ranging from 800 to 1,200 m.     

ATOC AND OTHER FACTORS AFFECTING THE DISTRIBUTION AND ABUNDANCE OF HUMPBACK WHALES (MEGAPTERA NOVAEANGLIAE) OFF THE NORTH SHORE OF KAUAI

Humpback whale diving behavior changes subtly when exposed to signals transmitted from the Acoustic Thermometry of Ocean Climate (ATOC) sound projector located 14 km offshore the island of Kauai. This study considered whether such responses would lead to changes in distribution and abundance. A land-based shore station measured humpback whale locations (scan samples) for both nearshore (<5 km) and offshore (5–10 km) areas. Control observations were made in 1994 and 1998. In 1998 multipleday blocks with ATOC transmissions were interspersed with multiple-day control blocks without transmissions. Sighting rates were higher in 1998 (ATOC) than in 1994 (control year), probably due to better sighting conditions, but may reflect increased population size. Sighting rates did not differ between control and ATOC conditions in 1998. A seasonal sighting peak was observed in both years. No vessel effect on sighting rate was detected in 1998.
There was no effect of ATOC on the distance from the shore station to whales, or the depth of water where pods were located. However, the distribution of whales shifted slightly eastward during the ATOC blocks and the mean distance between the ATOC source and pods was greater during transmissions. Nonetheless, more whales were found close to the source when it was on, suggesting a more variable response rather than simple avoidance, with whales found both closer to, and farther away from, the source during transmissions.     

FEEDING, SOCIAL AND MIGRATION BEHAVIOR OF BOWHEAD WHALES, BALAENA MYSTICETUS, IN BAFFIN BAY VS. THE BEAUFORT SEA—REGIONS WITH DIFFERENT AMOUNTS OF HUMAN ACTIVITY

This paper compares the behavior of bowhead whales of the Davis Strait/Baffin Bay stock, as observed along the east coast of Baffin Island in 1979–1986, with behavior of the Bering/Chukchi/Beaufort Sea stock observed in the Beaufort Sea in 1980–1986. All data used here were collected during late summer and early autumn in the absence of acute human disturbance. The behavioral repertoires of the two populations were similar. However, quantitative differences were found for whales engaged in all three activities studied: (1) Bowheads feeding in deep water off Isabella Bay, Baffin Island, had longer dives and surfacings, on average, than noted for bowheads feeding in the Beaufort Sea. (2) Among whales socializing in shallow water, we saw sexual interactions more often at Isabella Bay than in the Beaufort Sea. Calls emitted by socializing whales off Baffin Island were similar to those heard in the Chukchi and Beaufort Seas. However, pulsed tonal calls were longer off Baffin Island, and previously undescribed mechanical "crunch" sounds were recorded there near socializing bowheads. (3) During autumn migration, "fluke-out" dives were less common, and dive durations were longer, in the Beaufort Sea than off Baffin Island (P<0.001). Multivariate and other analyses indicated that some but not all differences can be ascribed to regional differences in the natural environment or in whale activities, However, during 1974–1986, Bering/Chukchi/Beaufort bowheads were exposed to more industrial, hunting and other human activity than Davis Strait/Baffin Bay bowheads. The "inconspicuous" behavior during autumn migration in the Beaufort may have been attributable to human activities, but causative links cannot be isolated.