MEASUREMENT OF PHOTOGRAPHIC QUALITY AND INDIVIDUAL DISTINCTIVENESS FOR THE PHOTOGRAPHIC IDENTIFICATION OF HUMPBACK WHALES, MEGAPTERA NOVAEANGLIAE

Accurate identification of humpback whales from photographic identification data depends on the quality of the photographs and the distinctiveness of the flukes. Criteria for evaluating photographic quality and individual distinctiveness were developed involving judgments about overall quality or distinctiveness and about specific aspects of each. These criteria were tested for the level of agreement among judges. The distinctiveness scheme was tested for the independence of distinctiveness judgments and photographic quality. Our results show that judges could agree when evaluating specific and overall aspects of photographic quality and individual distinctiveness. The level of agreement varied for different pairs of judges, and less adept judges were identified. Ability to agree on evaluations of photographic quality was independent of the experience of the judges. Overall photographic quality and overall distinctiveness were successfully predicted from more specific variables, but the agreement between judges for these was not significantly greater than the agreement for the overall measures judged directly. There was no correlation between individual distinctiveness and photographic quality for four of the five judges, but the power of this rest may be low. Analyses of photographic identification data frequently require evaluations of photographic quality and individual distinctiveness. To obtain reliable results from such analyses, evaluation schemes and judges should be tested to ensure reliable and consistent evaluations.     

TEMPORAL VARIABILITY IN FEATURES USED TO PHOTO-IDENTIFY HUMPBACK WHALES (MEGAPTERA NOVAEANGLIAE)

Photographs of 99 humpback whales of known age were analyzed to assess the temporal variability and recognizability of individually distinctive fluke and dorsal fin features used for photo-identification. Stable features tended to be morphological in nature (dorsal fin shape and edges, the trailing edge of the fluke, and the raised bumps on the caudal peduncle termed "knobs"). Transitory features typically were superficial marks (scarring, scratching, and pigmentation). The variability of several features was found to be age-dependent as well. Young animals sometimes experienced substantial change to their fluke coloration patterns, though this feature stabilized with age. Dorsal fin edges and fluke serration peaks tended to undergo more change in males than in females, with most changes occurring following sexual maturation. Peduncle knobs were the most stable feature, with no documented change in any age interval. Given the extreme stability (and hence recognizability) of dorsal fin shape and peduncle knobs, we suggest that photographs combining the dorsal fin and the caudal peduncle provide the most consistent way to reidentify humpback whales, particularly calves following weaning.     

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.     

REPRODUCTIVE RATES OF HUMPBACK WHALES OFF CALIFORNIA

From 1986 to 1996 we examined the reproductive rates, calving rates, and reproductive histories of mature females as part of photo-identification studies of humpback whales that feed off California, Oregon, and Washington during summer and fall. Annual reproductive rates were measured by two methods: proportion of all whales that were calves based on sightings (0.6%-5.9% per year, mean = 3.6%, SD = 1.6) and based on individually identified animals (1.1%-8.0% per year, mean = 4.1%, SD = 1.8). The reproductive rate based on sightings varied significantly by year ( G test, P < 0.001), region ( G test, P < 0.001), and by month ( G test, P < 0.05). Seventy-nine sexually mature females were identified with 97 calves out of a total of 844 known individuals over the 11-yr study. Mother-calf separation on the feeding grounds was recorded in several instances. The apparent reproductive rates of this population are considerably lower than rates of 4%–15% reported from other feeding areas for this species. Our estimates are likely biased downward because this population has been increasing at about 5% per year. Calves may have been missed due to early weaning and because of our sampling from small boats late in the season. We also found evidence of geographic segregation of mother-calf pairs within our large study area. Despite these factors, we conclude the reproductive rate of this population appears to be lower than has been reported in other areas.     

AN OCEAN-BASIN-WIDE MARK-RECAPTURE STUDY OF THE NORTH ATLANTIC HUMPBACK WHALE (MEGAPTERA NOVAEANGLIAE)

Although much is known about the humpback whale, Megaptera novaeangliae, regional studies have been unable to answer several questions that are central to the conservation and management of this endangered species. To resolve uncertainties about population size, as well as the spatial and genetic structure of the humpback whale population in the North Atlantic, we conducted a two-year ocean-basin-wide photographic and biopsy study in 1992-1993. Photographic and skin-biopsy sampling was conducted of animals in feeding and breeding areas throughout most of the range of this species in the North Atlantic, from the West Indies breeding grounds through all known feeding areas as far north as arctic Norway. A standardized sampling protocol was designed to maximize sample sizes while attempting to ensure equal probability of sampling, so that estimates of abundance would be as accurate and as precise as possible. During 666 d at sea aboard 28 vessels, 4,207 tail fluke photographs and 2,326 skin biopsies were collected. Molecular analyses of all biopsies included determination of sex, genotype using six microsatellite loci, and mitochondrial control region sequence. The photographs and microsatellite loci were used to identify 2,998 and 2,015 individual whales, respectively. Previously published results from this study have addressed spatial distribution, migration, and genetic relationships. Here, we present new estimates of total abundance in this ocean using photographic data, as well as overall and sex-specific estimates using biopsy data. We identify several potential sampling biases using only breeding-area samples and report a consistent mark-recapture estimate of oceanwide abundance derived from photographic identification, using both breeding and feeding-area data, of 10,600 (95% confidence interval 9,300-12,100). We also report a comparable, but less precise, biopsy-based estimate of 10,400 (95% confidence interval of 8,000-13,600). These estimates are significantly larger and more precise than estimates made for the 1980s, potentially reflecting population growth. In contrast, significantly lower and less consistent estimates were obtained using between-feeding-area or between-breeding-area sampling. Reasons for the lower estimates using the results of sampling in the same areas in subsequent years are discussed. Overall, the results of this ocean-basin-wide study demonstrate that an oceanwide approach to population assessment of baleen whales is practicable and results in a more comprehensive understanding of population abundance and biology than can be gained from smaller-scale efforts.     

SPATIAL AND SEASONAL DISTRIBUTION OF THE HUMPBACK WHALE, MEGAPTERA NOVAEANGLIAE, IN THE MEXICAN PACIFIC

From observations of the spatial distribution of humpback whales in the Mexican Pacific between 1981 and 1986, it is possible to recognize four subregions: 1) the southern coast of Baja California; 2) the northern Gulf of California, including the Midriff Islands; 3) the mainland coast of Mexico, including the Isla Isabel and Islas Tres Marias and 4) the Revillagigedo Archipelago. The seasonal distribution of whales near the Mexican mainland and the Revillagigedo Archipelago extends from November to May and is similar to that of other winter breeding grounds, including the Hawaiian Islands. Along the southern coast of Baja California, whales have been observed from September to April, possibly indicating a shorter migratory route. In the northern Gulf of California, however, humpback whales have been reported throughout the year and are occasionally observed feeding during both summer and winter months. The degree of individual movement between the four subregions is still unknown. The number of individual humpback whales identified photographically in recent years suggests that there ate more whales in the Mexican Pacific than previously reported.     

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.     

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.     

ABUNDANCE AND POPULATION TREND (1978-2001) OF WESTERN ARCTIC BOWHEAD WHALES SURVEYED NEAR BARROW, ALASKA

The 2001 survey of western Arctic (Bering, Chukchi, and Beaufort seas) bowhead whales was conducted from 5 April to 7 June near Barrow, Alaska. Visual observers recorded a total of 3,295 “new” (not seen before) and 532 “conditional” (possibly seen before) whales in 1,130 h of watch effort, including 121 new calves (3.7% of the new whales). Concurrent with the visual survey, passive acoustic surveillance was conducted almost continuously from 16 April to 31 May, resulting in 27,023 locations of vocalizing bowhead whales. The estimated number of whales within 4 km of the perch (N₄) was 7,025 (SE = 1,068). The estimated proportion of the whales within 4 km of the perch (P₄) was 0.862 (SE = 0.044, computed by a moving blocks bootstrap). Combining these, the abundance estimate (N₄/P₄) for 2001 is 10,470 (SE = 1, 351) with a 95% confidence interval of 8, 100–13, 500. The estimated annual rate of increase (ROI) of the population from 1978 to 2001 is 3.4% (95% CI 1.7%‐5%). Reports from hunters and results of an aerial survey in June 2001 indicate whales continued to pass Barrow after the survey had ended. In 2001 51% (572 h) of the watch was scored as occurring during “fair‐excellent” visibility conditions, somewhat lower than the average for all surveys since 1978. Sea ice in the leads and fog were the principle environmental factors affecting visibility for all years. The estimated rate of increase and the fact that the number of calves counted in 2001 is the highest ever recorded suggest a steady recovery of this population. Other populations of large balaenids, notably the North Atlantic right whale, have failed to recover despite 70 yr of protection. The recovery of the howhead whale is likely attributable to low anthropogenic mortality, a relatively pristine habitat, and a well‐managed subsistence hunt. Nonetheless, offshore oil development, increasing shipping traffic, changes in the Bering Sea ecosystem, sea ice retreat, and possibly killer whale predation within its range could impact this bowhead population and should be carefully monitored.     

RADIOTAGGING STUDIES OF BELUKHA WHALES (DELPHINAPTERUS LEUCAS) IN BRISTOL BAY, ALASKA

Research was conducted in Bristol Bay, Alaska, to determine the applicability of radiotagging to studies of behavior, distribution and movements of belukha whales. Backpack-style VHF transmitters were attached to two belukhas by pinning through the dorsal ridge. Both packages were shed after about 2 wk due to migration of the pin through the tissue. Movements of radio-tagged whales were essentially local within Kvichak Bay. Three basic respiration patterns were identified: surfacings that were grouped into breathing periods separated by longer dives; surfacings that did not occur during restricted breathing periods; and long-to-very-long surfacings separated by short-to-very-short dives. These patterns were interpreted as representing traveling, feeding and feeding or resting in very shallow water. Surface and dive interval data were used to calculate a correction factor of 2.75, which could then be applied to aerial survey counts to estimate the total number of belukha whales in the study area. Modifications to radio packages are necessary in order to increase retention time.     

AGE-LENGTH RELATIONSHIPS IN HUMPBACK WHALES: A COMPARISON OF STRANDINGS IN THE WESTERN NORTH ATLANTIC WITH COMMERCIAL CATCHES

Strandings of previously identified individuals, while rare, provide an opportunity to examine age-length relationships in humpback whales (Megaptera novacangliae) from the North Atlantic. Ages and lengths of 23 individuals are presented: 11 females and 12 males, 9 of known age and 14 with estimated minimum ages. Lengths ranged from 853 to 1, 430 cm, ages 0.5–17 yr. These individuals were generally smaller and more variable in size at age than reported from commercial catches. Fifteen of the stranded individuals were four years of age or younger, while few of the animals taken by whalers were this young, and these probably represented the larger individuals in these age categories. Thus the data presented herein help to give more definition to the early growth curve for the humpback whale than has previously been available. Growth equations illustrate a difference of about one meter in asymptotic length through age five between stranding and catch data. The close fit of growth models to data from younger and older animals separately and the difficulty of fitting a single growth model to animals of all ages, could indicate that a dynamic or staged growth pattern exists in this species.     

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.     

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.     

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.     

DISPERSAL, ROOKERY FIDELITY, AND METAPOPULATION STRUCTURE OF STELLER SEA LIONS (EUMETOPIAS JUBATUS) IN AN INCREASING AND A DECREASING POPULATION IN ALASKA

Over the past 24 yr, 8,596 Steller sea lion ( Eumetopias jubatus ) pups were branded on their natal rookeries throughout Alaska with the objectives of determining survival rates, recruitment, movements, and site fidelity. Our objectives here were to examine the extent of dispersal of Steller sea lions away from their natal rookeries, movements between stocks, and degree of natal rookery fidelity. Pups (<1 yr old) usually remained within 500 km of their natal rookery. Branded juveniles dispersed widely and were resighted at distances up to 1,785 km from their natal rookeries. Adults generally remained within 500 km of their natal rookeries. No interchange of breeding animals between the ES (eastern stock) and WS (western stock) was observed. Although natal rookery fidelity was prevalent, 33% of the 12 observations of females branded in the WS during 1987–1988 and 19% of the 29 observations of females branded in the ES during 1994–1995 were observed with newly born pups at sites other than their natal rookeries. Steller sea lions generally conformed to the metapopulation concept as depicted by Hanski and Simberloff (1997), with local breeding populations (rookeries) and movements among these local populations having the potential of affecting local dynamics.