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.

Samples were collected from 407 fin whales, Balaenoptera physalus, at four North Atlantic and one Mediterranean Sea summer feeding area as well as the Sea of Cortez in the Pacific Ocean. For each sample, the sex, the sequence of the first 288 nucleotides of the mitochondrial (mt) control region and the genotype at six microsatellite loci were determined. A significant degree of divergence was detected at all nuclear and mt loci between North Atlantic/Mediterranean Sea and the Sea of Cortez. However, the divergence time estimated from the mt sequences was substantially lower than the time elapsed since the rise of the Panama Isthmus, suggesting occasional gene flow between the North Pacific and North Atlantic ocean after the separation of the two oceans. Within the North Atlantic and Mediterranean Sea, significant levels of heterogeneity were observed in the mtDNA between the Mediterranean Sea, the eastern (Spain) and the western (the Gulf of Maine and the Gulf of St Lawrence) North Atlantic. Samples collected off West Greenland and Iceland could not be unequivocally assigned to either of the two areas. The homogeneity tests performed using the nuclear data revealed significant levels of divergence only between the Mediterranean Sea and the Gulf of St Lawrence or West Greenland. In conclusion, our results suggest the existence of several recently diverged populations in the North Atlantic and Mediterranean Sea, possibly with some limited gene flow between adjacent populations, a population structure which is consistent with earlier population models proposed by Kellogg, Ingebrigtsen, and Sergeant.

Samples from 136 humpback whales Megaptera novaeangliae, representing 5 feeding aggregations in the North Atlantic and 1 in the Antarctic, were analyzed with respect to the sequence variation in the mitochondria1 (mt) control region. A total of 288 base pairs was sequenced by direct sequencing of asymmetrically amplified DNA. Thirty-one different haplotypes were identified. The nucleotide diversity for the total sample was estimated to be 2.6 %, w h c h is high relative to other North Atlantic cetaceans. The degree of genetic differentiation in various subsets of the samples was estimated and tested for statistical significance by Monte Carlo simulations. Significant degrees of heterogeneity were found between the Antarctic and all North Atlantic areas, as well as between Iceland and the western North Atlantic samples. A genealogical tree was estimated for the 31 haplotypes and rooted with the homologous sequence from a fin whale Balaenoptera physalus. The branching pattern in the genealogical tree suggests that the North Atlantic Ocean has been populated by 2 independent influxes of humpback whales. The combined results from the homogeneity tests and the genealogical tree indicate that behaviour (in this case maternally directed site fidelity to a foraging area) can influence the population structure of marine cetaceans on an evolutionary time scale.

The consumption of fish, cephalopods and planktonic crustaceans by cetaceans in Icelandic and adjacent waters was estimated. The estimates were based on (i) abundance estimates from recent sighting surveys 89,95); (ii) seasonal variation in abund a n c e e s t i m a t e d f r o m s i g h t i n g s a n d / o r c a t c h d a t a f r o m w h a l i n g v e s s e l s ; a n d (iii) consumption rates calculated from the estimated biomass of cetaceans in the area throughout the year. A large number of assumptions had to be made, and these calculations were mainly intended to give an idea of the possible magnitude of consumption as well as to be a guidance for further research in this field. The total food consumption was estimated as 6.3 million tons in a smaller area defined as Icelandic and adjacent waters, and 8.8 million tons in the larger area north of 60 o N. Considering total consumption, fin whales (Balaenoptera physalus) and minke whales (B. acutorostrata) were the largest consumers in the area, followed by long-finned pilot whales (Globicephala melas) and northern bottlenose whales (Hyperoodon ampullatus). Crustaceans comprise around 50% of the total consumption within the study area while finfish and cephalopods comprised 27% and 22%, respectively, in the large area. Minke whales were the largest consumers of finfish, consuming more than 1 million tons of fish in Icelandic and adjacent waters.