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.
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.
Global warming poses particular challenges to migratory species, which face changes to the multiple environments occupied during migration. For many species, the timing of migration between summer and winter grounds and also within-season movements are crucial to maximise exploitation of temporarily abundant prey resources in feeding areas, themselves adapting to the warming planet. We investigated the temporal variation in the occurrence of fin (Balaenoptera physalus) and humpback whales (Megaptera novaeangliae) in a North Atlantic summer feeding ground, the Gulf of St. Lawrence (Canada), from 1984 to 2010 using a long-term study of individually identifiable animals. These two sympatric species both shifted their date of arrival at a previously undocumented rate of more than 1day per year earlier over the study period thus maintaining the approximate 2-week difference in arrival of the two species and enabling the maintenance of temporal niche separation. However, the departure date of both species also shifted earlier but at different rates resulting in increasing temporal overlap over the study period indicating that this separation may be starting to erode. Our analysis revealed that the trend in arrival was strongly related to earlier ice break-up and rising sea surface temperature, likely triggering earlier primary production. The observed changes in phenology in response to ocean warming are a remarkable example of phenotypic plasticity and may partly explain how baleen whales were able to survive a number of changes in climate over the last several million years. However, it is questionable whether the observed rate of change in timing can be maintained. Substantial modification to the distribution or annual life cycle of these species might be required to keep up with the ongoing warming of the oceans.
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.
Ecologically similar species may coexist when resource partitioning over time and space reduces interspecific competition. Understanding resource use within these species assemblages may help predict how species relative abundance might influence ecosystem functioning. In the Gulf of St. Lawrence, Canada, 4 species of rorqual whales (blue Balaenoptera musculus, fin B. physalus, minke B. acutorostrata and humpback Megaptera novaeangliae) coexist during the summer feeding period. They can be observed within hundreds of meters of one another, suggesting an overlap in ecological niches; yet fine-scale habitat use analyses suggest some resource partitioning. While major ecological changes have been observed in marine ecosystems, including the Gulf of St. Lawrence, we have little understanding of how the removal of predatory fish might cascade through ecosystems. Here, we take advantage of a 19 yr tissue collection subsequent to a fishery collapse (which occurred in 1992) to investigate trophic niche partitioning within a guild of rorqual whales following the loss of a key ecosystem component, groundfish. We analyzed stable isotope ratios for 626 rorqual individuals sampled between 1992 and 2010. Using Bayesian isotopic mixing models, we demonstrated that the 4 rorqual species segregated trophically by consuming different proportions of shared prey. An overall increase in δ 15 N values over the study period (post groundfish collapse), particularly for fin and humpback whales, suggested a progressive use of higher-trophic level prey, such as small pelagic fish, whereas the stability of blue whale diet over time confirmed their specialized feeding behaviour. This study provides the first longterm assessment of trophic ecology among rorqual populations on this Northwest Atlantic feeding ground, and evidence for differential resource use among large marine predators following ecosystem change.
Population spatial structuring among North Atlantic humpback whales Megaptera novaeangliae on the summer feeding grounds was investigated using movement patterns of identified individuals. We analysed the results from an intensive 2-year ocean-basin-scale investigation resulting in 1658 individuals identified by natural markings and 751 individuals by genetic markers supplemented with data from a long-term collaborative study with 3063 individuals identified by natural markings. Re-sighting distances ranged from o1 km to 42200 km. The frequencies (F) of resighting distances (D) observed in consecutive years were best modelled by an inverse allometric function (F=6631D À1.24 , r 2 = 0.984), reflecting high levels of site fidelity (median re-sighting distance o40 km) with occasional long-distance movement (5% of re-sightings 4550 km). The distribution of re-sighting distances differed east and west of 451W, with more long-distance movement in the east. This difference is consistent with regional patterns of prey distribution and predictability. Four feeding aggregations were identified: the Gulf of Maine, eastern Canada, West Greenland and the eastern North Atlantic. There was an exchange rate of 0.98% between the western feeding aggregations. The prevalence of long-distance movement in the east made delineation of possible additional feeding aggregations less clear. Limited exchange between sites separated by as little as tens of kilometres produced lower-level structuring within all feeding aggregations. Regional and temporal differences in movement patterns reflected similar foraging responses to varying patterns of prey availability and predictability. A negative relationship was shown between relative abundance of herring and sand lance in the Gulf of Maine and humpback whale movement from the Gulf of Maine to eastern Canada.
Understanding the factors influencing habitat selection is critical to improving management and conservation plans for large whales. Many studies have linked the distribution of cetaceans to basic environmental features such as underwater topography and sea surface temperature (SST), but the mechanisms underlying these relationships are poorly understood. Dynamic mesoscale processes like thermal fronts are prime candidates to link physiographic factors to whale distribution because they increase biological productivity and aggregate prey. However, previous studies of large whales have found little evidence of such associations, possibly because they were not at the appropriate spatio-temporal scales. We quantified the relationship between SST fronts and the distribution of blue Balaenoptera musculus, finback B. physalus, humpback Megaptera novaeangliae and minke B. acutorostrata whales in the northern Gulf of St. Lawrence. We compared the distribution of 1094 whale sightings collected from boat surveys conducted in 1996 to 2000 to the locations of frontal areas determined from 61 satellite maps. The distributions of whales and thermal fronts were highly correlated (random resampling and Mantel tests of matrix similarity). Spatial distributions differed among species, probably reflecting differences in feeding strategies. Identification of surface fronts from satellite imagery thus effectively complemented field observations of whales. These findings significantly increase our understanding of habitat quality in rorqual whales, and encourage a greater use of dynamic environmental variables in future studies of whale habitat use.KEY WORDS: Sea surface temperature · Thermal fronts · Blue whale · Finback whale · Humpback whale · Minke whale · Habitat selection · Gulf of St. Lawrence 335: 207-216, 2007 iness of krill (Murase et al. 2002) and fishes (Whitehead & Carscadden 1985). Patchiness of organisms in coastal ecosystems is often caused by the dynamic features of mesoscale oceanographic processes like fronts, eddies and upwellings (Olson & Backus 1985). These processes usually involve spatial scales of 1 to 10 km and temporal scales of 1 to 10 d (Hofmann & Powell 1998). In most cases, when these upwellings reach the surface, one of their manifestations is a thermal gradient between warm surface waters and cold upwelled waters. Such mesoscale oceanographic processes increase biological productivity and aggregate prey species (Olson & Backus 1985), thus influencing the distribution of several pelagic fish species (e.g. Fiedler & Bernard 1987, Podesta et al. 1993. Gaskin (1987) predicted that these transition zones between tidally mixed and thermally stratified areas could be an important feature of right whale habitat in the lower Bay of Fundy. This hypothesis was supported by some anecdotal evidence (Murison & Gaskin 1989) but not by quantitative results (Woodley & Gaskin 1996). Similarly, Baumgartner et al. (2003) suggested that spatial and interannual variability in right whale occurrence on the Sco...
To assess large‐whale stocks following the cessation of land‐based South Georgia whaling in 1965, we report three independent sighting databases: a cruise in 1997, observations from Bird Island (NW of South Georgia) between 1979 and 1998, and mariner sightings between 1992 and 1997. All species were rare, with sightings of southern right whales being the most common event. Two right whales photographed off South Georgia matched animals known from Peninsula Valdés, Argentina, a population known to be growing at 7%per annum. In contrast, blue and fin whales appeared to be less abundant. A single blue whale mother‐calf pair was observed off the Shag Rocks in February 1997. Extirpation of animals from this particular feeding ground is the most likely reason for ongoing low numbers of all species. Other factors may include competition for krill by traditional predators such as penguins and seals and more recently by humans, an unusually high rate of natural mortality, habitat change such as alteration in sea ice coverage, and/or the impact of ongoing whaling. The history of this critical area of large‐whale habitat and this report demonstrate the need for improved, consistent longterm monitoring of population trends for these depleted stocks.
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