Populations of seals, sea lions, and sea otters have sequentially collapsed over large areas of the northern North Pacific Ocean and southern Bering Sea during the last several decades. A bottom-up nutritional limitation mechanism induced by physical oceanographic change or competition with fisheries was long thought to be largely responsible for these declines. The current weight of evidence is more consistent with top-down forcing. Increased predation by killer whales probably drove the sea otter collapse and may have been responsible for the earlier pinniped declines as well. We propose that decimation of the great whales by postWorld War II industrial whaling caused the great whales' foremost natural predators, killer whales, to begin feeding more intensively on the smaller marine mammals, thus ''fishing-down'' this element of the marine food web. The timing of these events, information on the abundance, diet, and foraging behavior of both predators and prey, and feasibility analyses based on demographic and energetic modeling are all consistent with this hypothesis.T he abrupt decline of the western stock of Steller sea lions (Eumetopias jubatus) h across most of the northern North Pacific Ocean and southern Bering Sea is one of the world's most well known yet poorly understood marine conservation problems. For years, scientists attributed this decline to nutritional limitation, the presumed consequence of a climate regime shift and蜑or competition with regional fisheries (1). Although fisheries and regime shifts undoubtedly influenced both the fishes and their associated food webs (2-5), several recent reviews of the available information on sea lions and their environment, including an assessment by the National Research Council, cast doubt on the nutritional limitation hypothesis (6, 7), notwithstanding evidence from field and laboratory studies that diet quality is a factor in sea lion energetics (8). The doubt stems from three main findings. First, most measures of behavior, physiology, and morphology from surviving adult sea lions and pups in the western Gulf of Alaska and Aleutian Islands are inconsistent with nutritional limitation. These animals have better body condition, reduced foraging effort, and reduced field metabolic rates relative to similar measures from the increasing sea lion population in southeast Alaska (7). Second, sea lion prey is abundant in most areas of the decline (9). Known changes in prey availability and other features of the oceanic ecosystem are particularly incongruous with the most precipitous phase of the decline, which occurred during the mid-to late 1980s, and can be accounted for only by greatly increased adult mortality (6). Third, populations of piscivorous sea birds, many of which feed on earlier life stages of the same fish species consumed by sea lions, have remained stable or increased in the same area and over the same period that the sea lions have declined (10). Top-down forcing now appears to have been an important contributor to declines of Steller sea lion...
Cetacean-habitat modeling, although still in the early stages of development, represents a potentially powerful tool for predicting cetacean distributions and understanding the ecological processes determining these distributions. Marine ecosystems vary temporally on diel to decadal scales and spatially on scales from several meters to 1000s of kilometers. Many cetacean species are wideranging and respond to this variability by changes in distribution patterns. Cetacean-habitat models have already been used to incorporate this variability into management applications, including improvement of abundance estimates, development of marine protected areas, and understanding cetacean-fisheries interactions. We present a review of the development of cetacean-habitat models, organized according to the primary steps involved in the modeling process. Topics covered include purposes for which cetacean-habitat models are developed, scale issues in marine ecosystems, cetacean and habitat data collection, descriptive and statistical modeling techniques, model selection, and model evaluation. To date, descriptive statistical techniques have been used to explore cetacean-habitat relationships for selected species in specific areas; the numbers of species and geographic areas examined using computationally intensive statistic modeling techniques are considerably less, and the development of models to test specific hypotheses about the ecological processes determining cetacean distributions has just begun. Future directions in cetacean-habitat modeling span a wide range of possibilities, from development of basic modeling techniques to addressing important ecological questions.
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