After nearly a century of recovery from overhunting, sea otter populations are in abrupt decline over large areas of western Alaska. Increased killer whale predation is the likely cause of these declines. Elevated sea urchin density and the consequent deforestation of kelp beds in the nearshore community demonstrate that the otter's keystone role has been reduced or eliminated. This chain of interactions was probably initiated by anthropogenic changes in the offshore oceanic ecosystem.

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 and͞or 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...

Locomotor activity by diving marine mammals is accomplished while breath-holding and often exceeds predicted aerobic capacities. Video sequences of freely diving seals and whales wearing submersible cameras reveal a behavioral strategy that improves energetic efficiency in these animals. Prolonged gliding (greater than 78% descent duration) occurred during dives exceeding 80 meters in depth. Gliding was attributed to buoyancy changes with lung compression at depth. By modifying locomotor patterns to take advantage of these physical changes, Weddell seals realized a 9.2 to 59.6% reduction in diving energetic costs. This energy-conserving strategy allows marine mammals to increase aerobic dive duration and achieve remarkable depths despite limited oxygen availability when submerged.

The spatial scale at which organisms respond to human activity can affect both ecological function and conservation planning. Yet little is known regarding the spatial scale at which distinct behaviors related to reproduction and survival are impacted by human interference. Here we provide a novel approach to estimating the spatial scale at which a top predator, the puma (Puma concolor), responds to human development when it is moving, feeding, communicating, and denning. We find that reproductive behaviors (communication and denning) require at least a 4× larger buffer from human development than non-reproductive behaviors (movement and feeding). In addition, pumas give a wider berth to types of human development that provide a more consistent source of human interference (neighborhoods) than they do to those in which human presence is more intermittent (arterial roads with speeds >35 mph). Neighborhoods were a deterrent to pumas regardless of behavior, while arterial roads only deterred pumas when they were communicating and denning. Female pumas were less deterred by human development than males, but they showed larger variation in their responses overall. Our behaviorally explicit approach to modeling animal response to human activity can be used as a novel tool to assess habitat quality, identify wildlife corridors, and mitigate human-wildlife conflict.

Foraging by mammals is a complex suite of behaviors that can entail high energetic costs associated with supporting basal metabolism, locomotion and the digestion of prey. To determine the contribution of these various costs in a free-ranging marine mammal, we measured the post-dive oxygen consumption of adult Weddell seals (N=9) performing foraging and nonforaging dives from an isolated ice hole in McMurdo Sound, Antarctica. Dives were classified according to behavior as monitored by an attached video-data logging system (recording activity, time, depth, velocity and stroking). We found that recovery oxygen consumption showed a biphasic relationship with dive duration that corresponded to the onset of plasma lactate accumulation at approximately 23·min. Locomotor costs for diving Weddell seals increased linearly with the number of strokes taken according to the relationship: locomotor cost = -3.78+0.04 × stroke number (r 2 =0.74, N=90 dives), where locomotor cost is in ml·O2·kg -1 . Foraging dives in which seals ingested Pleuragramma antarcticum resulted in a 44.7% increase in recovery oxygen consumption compared to non-foraging dives, which we attributed to the digestion and warming of prey. The results show that the energy expended in digestion for a free-ranging marine mammal are additive to locomotor and basal costs. By accounting for each of these costs and monitoring stroking mechanics, it is possible to estimate the aerobic cost of diving in free-ranging seals where cryptic behavior and remote locations prevent direct energetic measurements.

Large body size, carnivory, and endothermic costs lead to exceptionally high caloric demands in many mammalian predators. The potential impact on prey resources may be marked but is difficult to demonstrate because of the mobility, sparseness, and cryptic nature of these animals. In this study, we developed a method based on comparative bioenergetics and demographic modeling to evaluate predator effects and then used this approach to assess the potential impact of killer whales on sea otter and Steller sea lion populations in the Aleutian Islands. Daily caloric requirements of killer whales determined from allometric regressions for field metabolic rate show that an adult killer whale requires 51-59 kcal·kg Ϫ1 ·d Ϫ1 (2.5-2.9 W/kg). Caloric values of prey items determined by bomb calorimetry ranged from 41 630 kcal for an adult female sea otter to sequentially higher values for male otters, sea lion pups, and adult Steller sea lions. Integrating these results with demographic changes in marine mammal populations show that fewer than 40 killer whales could have caused the recent Steller sea lion decline in the Aleutian archipelago; a pod of five individuals could account for the decline in sea otters and the continued suppression of sea lions. The collapse of the historical prey base of killer whales due to human whaling may have contributed to a sequential dietary switch from high to low caloric value prey, thereby initiating these declines. This study demonstrates that a combined physiological-demographic approach increases our ability to critically evaluate the potential impact of a predator on community structure and enables us to define underlying mechanisms that drive or constrain top-down forcing in dynamic ecosystems.

Mammals re-entered the oceans less than 60 million years ago. The transition from a terrestrial to an aquatic lifestyle required extreme morphological and behavioural modi¢cations concomitant with fundamentally di¡erent locomotor mechanisms for moving on land and through water. Energetic transport costs typically re£ect such di¡erent locomotor modes, but can not be discerned from the fossil record. In this study the energetic challenges associated with changing from terrestrial to aquatic locomotion in primitive marine mammals are examined by comparing the transport, maintenance and locomotor costs of extant mammals varying in degree of aquatic specialization. The results indicate that running and swimming specialists have converged on an energetic optimum for locomotion. An allometric expression, COT TOT 7.79 mass À0X29 (r 2 0.83, n 6 species), describes the total cost of transport in J kg À1 m À1 for swimming marine mammals ranging in size from 21kg to 15 000 kg. This relation is indistinguishable from that describing total transport costs in running mammals. In contrast, the transitional lifestyle of semi-aquatic mammals, similar to that of ancestral marine mammals, incurs costs that are 2.4^5.1 times higher than locomotor specialists. These patterns suggest that primitive marine mammals confronted an energetic hurdle before returning to costs reminiscent of their terrestrial ancestry, and may have reached an evolutionary limit for energetic optimization during swimming.

The hunting behavior of a marine mammal was studied beneath the Antarctic fast ice with an animal-borne video system and data recorder. Weddell seals stalked large Antarctic cod and the smaller subice fish Pagothenia borchgrevinki, often with the under-ice surface for backlighting, which implies that vision is important for hunting. They approached to within centimeters of cod without startling the fish. Seals flushed P. borchgrevinki by blowing air into subice crevices or pursued them into the platelet ice. These observations highlight the broad range of insights that are possible with simultaneous recordings of video, audio, three-dimensional dive paths, and locomotor effort.