Summary 1.The status of small cetaceans in the North Sea and adjacent waters has been of concern for many years. Shipboard and aerial line transect surveys were conducted to provide accurate and precise estimates of abundance as a basis for conservation strategy in European waters. 2. The survey, known as SCANS (Small Cetacean Abundance in the North Sea), was conducted in summer 1994 and designed to generate precise and unbiased abundance estimates. Thus the intensity of survey was high, and data collection and analysis methods allowed for the probability of detection of animals on the transect line being less than unity and, for shipboard surveys, also allowed for animal movement in response to the survey platform. 3. Shipboard transects covered 20 000 km in an area of 890 000 km 2 . Aerial transects covered 7000 km in an area of 150 000 km 2 . 4. Three species dominated the data. Harbour porpoise Phocoena phocoena were encountered throughout the survey area except in the Channel and the southern North Sea. Whitebeaked dolphin Lagenorhynchus albirostris and minke whale Balaenoptera acutorostrata were found mainly in the north-western North Sea. 5. Phocoena phocoena abundance for the entire survey area was estimated as 341 366 [coefficient of variation (CV) = 0·14; 95% confidence interval (CI) = 260 000-449 000]. The estimated number of B. acutorostrata was 8445 (CV = 0·24; 95% CI 5000-13 500). The estimate for L. albirostris based on confirmed sightings of this species was 7856 (CV = 0·30; 95% CI = 4000 -13 000). When Atlantic whitesided dolphin Lagenorhynchus acutus and Lagenorhynchus spp. sightings were included, this estimate increased to 11 760 (CV = 0·26; 95% CI 5900-18 500). 6. Shortbeaked common dolphin Delphinus delphis were found almost exclusively in the Celtic Sea. Abundance was estimated as 75 450 (CV = 0·67; 95% CI = 23 000 -149 000). 7. Current assessments and recommendations by international fora concerning the impact on P. phocoena of bycatch in gillnet fisheries in the North Sea and adjacent waters are based on these estimates.
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.
Population genetic structure of North Atlantic killer whale samples was resolved from differences in allele frequencies of 17 microsatellite loci, mtDNA control region haplotype frequencies and for a subset of samples, using complete mitogenome sequences. Three significantly differentiated populations were identified. Differentiation based on microsatellite allele frequencies was greater between the two allopatric populations than between the two pairs of partially sympatric populations. Spatial clustering of individuals within each of these populations overlaps with the distribution of particular prey resources: herring, mackerel and tuna, which each population has been seen predating. Phylogenetic analyses using complete mitogenomes suggested two populations could have resulted from single founding events and subsequent matrilineal expansion. The third population, which was sampled at lower latitudes and lower density, consisted of maternal lineages from three highly divergent clades. Pairwise population differentiation was greater for estimates based on mtDNA control region haplotype frequencies than for estimates based on microsatellite allele frequencies, and there were no mitogenome haplotypes shared among populations. This suggests low or no female migration and that gene flow was primarily male mediated when populations spatially and temporally overlap. These results demonstrate that genetic differentiation can arise through resource specialization in the absence of physical barriers to gene flow.
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