We report on a wintering area off the Pacific coast of Central America for humpback whales (Megaptera novaeangliae) migrating from feeding areas off Antarctica. We document seven individuals, including a mother/calf pair, that made this migration (approx. 8300km), the longest movement undertaken by any mammal. Whales were observed as far north as 11 degrees N off Costa Rica, in an area also used by a boreal population during the opposite winter season, resulting in unique spatial overlap between Northern and Southern Hemisphere populations. The occurrence of such a northerly wintering area is coincident with the development of an equatorial tongue of cold water in the eastern South Pacific, a pattern that is repeated in the eastern South Atlantic. A survey of location and water temperature at the wintering areas worldwide indicates that they are found in warm waters (21.1-28.3 degrees C), irrespective of latitude. We contend that while availability of suitable reproductive habitat in the wintering areas is important at the fine scale, water temperature influences whale distribution at the basin scale. Calf development in warm water may lead to larger adult size and increased reproductive success, a strategy that supports the energy conservation hypothesis as a reason for migration.
The ability to recognize individual animals has substantially increased our knowledge of the biology and behaviour of many taxa. However, not all species lend themselves to this approach, either because of insufficient phenotypic variation or because tag attachment is not feasible. The use of genetic markers ('tags') represents a viable alternative to traditional methods of individual recognition, as they are permanent and exist in all individuals. We tested the use of genetic markers as the primary means of identifying individuals in a study of humpback whales in the North Atlantic Ocean. Analysis of six microsatellite loci among 3,060 skin samples collected throughout this ocean allowed the unequivocal identification of individuals. Analysis of 692 'recaptures', identified by their genotype, revealed individual local and migratory movements of up to 10,000 km, limited exchange among summer feeding grounds, and mixing in winter breeding areas, and also allowed the first estimates of animal abundance based solely on genotypic data. Our study demonstrates that genetic tagging is not only feasible, but generates data (for example, on sex) that can be valuable when interpreting the results of tagging experiments.
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 mitochondria1 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 Present address: SMITH E T A L . : MARK-RECAPTURE STUDY 3 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.
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.
Certain populations of killer whales Orcinus orca feed primarily or exclusively on marine mammals. However, whether or not baleen whales represent an important prey source for killer whales is debatable. A hypothesis by Springer et al. (2003) suggested that overexploitation of large whales by industrial whaling forced killer whales to prey-switch from baleen whales to pinnipeds and sea otters, resulting in population declines for these smaller marine mammals in the North Pacific and southern Bering Sea. This prey-switching hypothesis is in part contingent upon the idea that killer whales commonly attack mysticetes while they are in these high-latitude areas. In this study, we used photographic and sighting data from long-term studies of baleen whales in 24 regions worldwide to determine the proportion of whales that bear scars (rake marks) from killer whale attacks, and to examine the timing of scar acquisition. The results of this study show that there is considerable geographic variation in the proportion of whales with rake marks, ranging from 0% to > 40% in different regions. In every region, the great majority of the scars seen were present on the whales' bodies when the animals were first sighted. Less than 7% (9 of 132) of scarred humpback whales with multi-year sighting histories acquired new scars after the first sighting. This suggests that most killer whale attacks on baleen whales target young animals, probably calves on their first migration from low-latitude breeding and calving areas to high-latitude feeding grounds. Overall, our results imply that adult baleen whales are not an important prey source for killer whales in high latitudes, and therefore that one of the primary assumptions underlying the Springer et al. (2003) prey-switching hypothesis (and its purported link to industrial whaling) is invalid.
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