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.     

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.     

ACOUSTIC DETECTIONS OF SINGING HUMPBACK WHALES IN DEEP WATERS OFF THE BRITISH ISLES

From October 1996 through September 1998, we used bottom-mounted hydrophone arrays to monitor deep-water areas north and west of the British Isles for songs of humpback whales ( Megaptera novaeangliae ). Singing humpbacks were consistently detected between October and March from the Shetland-Faroe Islands south to waters west of the English Channel. Temporal and geographic patterns of song detections, and movements of individually tracked whales, exhibited a southwesterly trend over this period, but with no corresponding northward trend between April and September. These results, together with a review of historical data from this area, suggest that the offshore waters of the British Isles represent a migration corridor for humpbacks, at least some of which summer in Norwegian (and possibly eastern Icelandic) waters. The migratory destination of the detected animals remains unknown, but the limited data suggest that these whales are bound primarily for the West Indies rather than historical breeding areas off the northwestern coast of Africa. Humpbacks detected in British waters after early to mid-March probably do not undertake a full migration to the tropics. These data provide further evidence that singing is not confined to tropical waters in winter, but occurs commonly on migration even in high latitudes.     

SEASONAL AND DIURNAL TRENDS OF CHORUSING HUMPBACK WHALES WINTERING IN WATERS OFF WESTERN MAUI

A portable data logger controlled by a Tattletale 7 microcontroller was used to record humpback whale choruses during the 1998 humpback whale winter season in Hawaii. The data logger sampled the sounds for four minutes every half hour using a digitizing rate of 2 kHz, and the data were stored on a hard disk. The results between January and April showed a peak in the sound pressure level between mid-February and mid-March. This peak of approximately 120 dB re 1 μPa coincided with the peak in the number of whales sighted by aerial survey on 7 March 1998. The choruses had spectral peaks at 315 Hz and 630 Hz. Some of the sounds at 630 Hz were second harmonics of the 315 Hz peak and others were not. The data also indicated a diurnal pattern in the sound pressure level, with levels at night significantly louder than the daytime levels. The sound levels began to increase during sunset and remained relatively high until sunrise, when they progressively decreased to a minimum. The nighttime peak occurred within an hour before and after midnight, and the daytime minimum occurred between 1100 and 1500. That more humpback whales appear to sing at night may reflect a switch to sexual advertisement as the primary male mating strategy at this time. It may also indicate that daylight and vision play key roles in the formation of competitive groups. It is suggested that the relative number of humpback whales in a given locale may be estimated by monitoring changes in sound pressure levels.     

MITOCHONDRIAL DNA VARIATION AND MATERNAL GENE FLOW AMONG HUMPBACK WHALES OF THE SOUTHERN HEMISPHERE

Samples of skin tissue were collected by biopsy darting from humpback whales ( Megaptera novaeangliae ) in six seasonal habitats representing three stocks and four regions: Groups IV (western Australia), V western component (eastern Australia), V eastern component (New Zealand and Tonga) and VI (the Antarctic Peninsula and Gorgona Island, Colombia, South America) of the Southern Hemisphere. A variable section of the mitochondrial DNA control region was amplified and sequenced from 84 of these individuals, distinguishing a total of 48 unique sequences ( i. e. , mtDNA nucleotypes). Phylogenetic reconstructions suggested that these nucleotypes form three clades, corresponding to those previously described in a world-wide survey of humpback whale mtDNA variation, although bootstrap support for two of the clades was relatively low (<50%). An analysis of variance adapted for molecular information showed significant differentiation of nucleotypes among the three Groups (Stocks) and heterogeneity of haplotype diversity among the four regions. A pattern of interchange within and between oceanic basins was demonstrated by the presence of shared identical nucleotypes among humpback whales in regions of the Southern and Northern Hemispheres.     

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.     

REACTIONS OF HUMPBACK WHALES TO SKIN BIOPSY SAMPLING ON A WEST INDIES BREEDING GROUND

Abstract: Reactions of humpback whales, Megaptera novaeangliae , to the taking of skin biopsies and to associated activities were studied on one of their principal West Indies breeding grounds on Silver Bank (Dominican Republic). Results were in some cases different from those reported from a similar study of this species in a high-latitude feeding area. Almost half (44.1%) of 565 biopsied whales showed no immediate reaction to a hit, while a further 22.5% showed only low-level reactions. A total of 375 (87.8%) of 427 misses involved no reaction. Only one strong reaction was recorded. Behavior changes were recorded following 31 (5.5%) of 569 hits, and 18 (4.5%) of 404 misses. Evasive behavior related to vessel approach was exhibited prior to 72 (12.0%) of 598 hits and 100 (24.1%) of 415 misses. Mothers showed significantly fewer reactions to hits than other whales, and a similar frequency and type of behavior changes, although they tended to be more evasive before a shot was made. Presumed males in competitive groups also showed significantly fewer reactions to shots, and very few behavior changes. Overall, this study supports the belief that the biopsy itself has little effect on a whale and that, if the associated vessel approach is conducted with care, samples can usually be taken with minimal disturbance to the target animal. However, approaches may affect the probability of obtaining fluke photographs for individual identification.     

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.     

ASSESSMENT OF HETEROGENEITY IN SIGHTING PROBABILITIES OF HUMPBACK WHALES WITHIN VIEWING RANGE OF CAPE VIDAL, SOUTH AFRICA

The migrations of humpback whales on the African east coast were monitored between 1988 and 1991 at Cape Vidal, South Africa. Shore-based surveys of the northward migration were undertaken each winter, and a survey of the southward migration was undertaken in 1990, from an approximately 60-m-high vantage point on a headland. Independent-observer surveys were carried out in both 1990 (22 d) and 1991 (51 d) to determine the proportion of the population within the survey area that were being missed by observers using a single mark-release model. Results were stratified into three distance intervals from the shore and three sighting condition intervals; there were constant sighting probabilities from the south tower under different sighting conditions, while those from the north tower increased slightly as sighting conditions improved. Sighting probabilities from both towers were highest in the intermediate distance interval and decreased in both the inshore and offshore regions.     

BEHAVIORAL RESPONSES OF EAST AUSTRALIAN HUMPBACK WHALES MEGAPTERA NOVAEANGLIAE TO BIOPSY SAMPLING

The response of migrating humpback whales to biopsy sampling was investigated off North Stradbroke Island, South East Queensland. Whales were allocated a behavioral category prior to biopsy sampling according to the general behavior of their pod. Behavioral reactions were recorded after each attempt. Sex was determined using a molecular genetic technique.
Detectable reactions occurred in 41.6% of successful biopsy attempts, a significantly lower response rate than that reported by two studies carried out on the feeding and breeding grounds of the North Atlantic. There was no difference in the response rate of whales on their northward or southward migration. Pod size was not an important factor in predicting the response of an individual. Females responded to biopsy sampling at a significantly higher rate than males.
Our results indicate that a substantial difference in response rate can occur between studies. Factors such as the type of boat used and the prior exposure of whales to human impact may be of importance. Our study suggests that female humpback whales may be particularly responsive to human disturbances. Overall, however, biopsy sampling has minimal impact on humpback whales.     

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.     

ESTIMATES OF THE NUMBERS OF HUMPBACK WHALES OBSERVED MIGRATING PAST CAPE VIDAL, SOUTH AFRICA, 1988–1991

A recovery-monitoring program of the African east coast humpback whale population was carried out through shore-based visual surveys from Cape Vidal, northern Natal. Surveys of the northward migration were undertaken each winter from 1988 to 1991, and a survey of the southward migration was undertaken in 1990. Independent observer surveys were undertaken during June 1990 and during the entire 1991 survey. Hourly densities of groups sighted each day were adjusted for groups missed by observers with distance from the shore and under different sighting conditions. Densities were multiplied by 24 h and the mean group size of the survey year to give resulting daily densities of individuals, which were summed to provide totals of whales sighted during each year's survey. The best estimate of population size was 1,711 (made during the northward migration of 1990), although this is likely to be biased downwards by a proportion of the population passing outside of observers'view. Bootstrapping of the 1991 daily data resulted in CVs between 11.4% and 12.2%. The numbers sighted show the population to have undergone considerable recovery since protection in October 1963.     

FISHING GEAR INVOLVED IN ENTANGLEMENTS OF RIGHT AND HUMPBACK WHALES

Interactions between marine mammals and fishing gear are an issue of global concern. Entanglements in the western North Atlantic are a major source of injury and mortality for endangered large whales. In this study, entanglements of 31 right whales ( Eubalaena glacialis ) and 30 humpback whales ( Megaptera novaeangliae ) were analyzed to determine the types and parts of gear involved. When gear was identified, 89% ( n = 32) of the entanglements were attributed to pot and gill net gear; however, a wide range of specific gear types were implicated. Despite gear recovery, gear type was not identified in 20% ( n = 9) of the cases. Although pot gear was recovered from both species equally, gill net gear was less frequently retrieved from right whales ( n = 2) than humpback whales ( n = 11). When gear part was identified, 81% ( n = 21) involved entanglements in buoy line and/or groundline. For right whales, the most common point of gear attachment was the mouth (77.4%); for humpback whales, the tail (53%) and the mouth (43%) were common attachment sites. Four right and three humpback whales in this sample were known to have died subsequent to entanglement. However, when identified, the gear types and parts involved in lethal cases were not substantially different from entanglements with non-lethal outcomes. Large whales can become entangled in a wide variety of fishing gear types and parts, and additional insight will depend on continued efforts to document entanglements and recover associated gear.     

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.     

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.     

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.     

NORTH PACIFIC HUMPBACK WHALE SONGS: A COMPARISON OF SOUTHEAST ALASKAN FEEDING GROUND SONGS WITH HAWAIIAN WINTERING GROUND SONGS

Humpback whales sing long, complex songs on their wintering grounds. On 25 August 1979 and 3 September 1981, we made recordings of humpback whale songs in southeastern Alaska, showing that humpback whales also sing on the summer feeding grounds. Both these Alaskan samples are songs in that they are repeating cyclical sound patterns and follow the known structure for humpback whale song. The Alaskan songs contain all the same material sung in the same order as that heard off Mexico and Hawaii in the surrounding wintering seasons. However, song, theme and some phrase durations are abbreviated in the Alaskan songs. The recording of these two songs represents the full sample of song recorded from 155 days over five years of attempting to record humpback whale song in Alaskan waters.     

POPULATION CHARACTERISTICS AND MIGRATION OF SUMMER AND LATE-SEASON HUMPBACK WHALES (MEGAPTERA NOVAEANGLIAE) IN SOUTHEASTERN ALASKA

A total of 326 humpback whales (Megaptera novaeangliae) were individually identified in southeastern Alaska during five summer seasons (July to September) and four late seasons (November to February) spanning the years 1979 to 1983. Peak numbers of whales were found late in August or early in September. Whales arrived 1–2 wk later in 1982 than in 1981. Whales sighted in both the summer and late seasons of 1981 and 1982 remained about 3.7 mo and one whale remained for at least 4.9 mo. Humpback whales from southeastern Alaska wintered in Hawaiian or Mexican waters, but generally did not travel to other feeding regions. The most rapid migratory transit between Hawaii and southeastern Alaska was 79 d. Based on mark-recapture analyses of the photographic data, we estimate a population of 270–372 whales in the southeastern Alaska feeding herd.     

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.     

INTERACTIONS OF SINGING HUMPBACK WHALES WITH OTHER MALES

Two non-mutually exclusive hypotheses on the function of the humpback whale song are: (1) it attracts females to the male singer; (2) it is a male-male display, that may order status. To evaluate these, from 24 January-13 April 1997 off Maui, Hawaii, 42 singers were located, audio-recorded, photo-identified and monitored for interactions with other whales. Whales that joined singers were biopsy sampled for molecular determination of sex. In 76% (32 of 42) of the interactions, a lone non-singing adult joined the singer. In the remainder, singers stopped singing and joined a nearby group or accompanied other whales. In 81% (26 of 32) instances where a lone adult joined a singer, the pair split again within minutes; otherwise a group formed. In one such group the pair became a trio and eventually joined a competitive group. Behavior in joining/splitting interactions ranged from a single pass-by, to surface activity such as tail lobs and breaches. The sex of 22 joiners was determined: 14 genetically and eight behaviorally. All were males. Humpback whale song preceded, and at times followed, male-male interactions of variable duration and agonistic level in and around the breeding season. If considered within the context of a proposed dominance polygyny mating system, these observations appear to support speculation that the song may function in male social ordering.