Many marine mammal populations are currently recovering from population depletion after overharvest. As marine mammals are often important predators in shaping marine ecosystems, there is a need to understand the impacts of recovering populations on other species and the marine ecosystem as a whole. The depletion and subsequent recovery of these species presents biologists with natural experiments to study their ecology, including drivers of their population dynamics and the function of the species in the ecosystem. This dissertation focuses on the recovery of a translocated population of sea otters (Enhydra lutris kenyoni) in Washington State. The presence or absence of sea otters, a keystone species, can dramatically influence marine community structure. The overall aim of this dissertation was to utilize the natural experiment of sea otter translocation to Washington State to understand drivers of sea otter population dynamics as well as the ecological role that sea otters play in Washington State. In Chapter 2, my coauthors and I found that the sea otter population in Washington has grown from an estimated 21 adult sea otters in 1977 to 2,336 adult sea otters in 2019, and the population is predicted to continue to grow and expand primarily to the south of the current range over the next 25 years. We also estimated that Washington State can support twice as many sea otters than previously estimated (equilibrium abundance of 6,080 vs. 2,734 sea otters), and that estimates of mean equilibrium density in currently occupied areas had the largest impact on predictions of population growth and range expansion. In Chapter 3, we quantified how sea otter population status (i.e., sea otter cumulative density) and habitat type (i.e., sea otter foraging in open water, kelp canopy, emergent rock, or intertidal) influence sea otter diet, and found that habitat was 1.77 times more important than sea otter population status in determining sea otter diet composition. We also found that sea otter long-term average rate of energy intake and diet diversity were negatively and positively correlated with sea otter cumulative density, respectively. In Chapter 4, we demonstrated the ecological role of sea otters in the nearshore marine ecosystem in Washington as a keystone species. We found that temporal transitions in the amount of kelp canopy were related to the duration of sea otter occupation, and that this relationship was more complex than a simple linear function. We also found that sea urchins were present at higher densities at sites more recently occupied by sea otters compared to long-occupied sites. In Chapter 5, we demonstrated the impact of sea otters as a recovering predator on the Pacific razor clam (Siliqua patula). We found that the magnitude of sea otter predation effects varied over time and space, with sea otter-caused razor clam mortality surpassing natural mortality in 2018 at Kalaloch Beach, occupied by sea otters since 2005. We also found that sea otters selectively consume the larger “recruit” size razor clams, the size that is also targeted in the recreational fishery, despite the smaller pre-recruit size clams being more abundant. Collectively, these results provide a deeper understanding of sea otter recolonization in Washington State as well as the ecological consequences of this recolonization.

The growing sea otter population in southern Southeast Alaska is impacting commercial shellfish, through foraging and expanding in range and abundance except where hunted for subsistence. Sea otters and their prey have coexisted in the North Pacific Ocean for approximately 750,000 years, but due to exploitation of sea otters from the 1770s until 1911, the species became extinct over much of its range, including southern Southeast Alaska. Subsequently, invertebrate species flourished and were commercially targeted in the late 1900s. Sea otters were relocated (n = 106) to southern Southeast Alaska in 1968. In this dissertation, I evaluated this marine mammal-fisheries conflict through multiple approaches. In Chapter 1, I analyzed geoduck clam and red sea urchin abundance surveys (1994-2012) and catch and effort data from commercial Dungeness crab fisheries (1969-2010) to identify interactions between sea otters and commercial shellfish. In Chapter 2, I collected geo-locations from 30 instrumented sea otters (2011-2014) to identify space use and range expansion. In Chapter 3, I collected sea otter abundance and distribution data from fixed wing aircraft (2010-2014) and observational forage data from sea otters (2010-2013) to determine contemporary population growth and consumption of commercially important shellfish by sea otters. The sea otter population in southern Southeast Alaska has grown from 106 to an estimated 13,139 individuals between 1968 and 2011 with an annual growth rate of 12% and expansion of its range by 117 km2 y-1. Results from a before-after, control-impact analysis indicate that sea otters are rapidly impacting red sea urchin and significantly reducing geoduck clam densities. Further, breakpoints predicted from regression models of Dungeness crab catch are correlated with known sea otter colonization timing. Forty-six percent of the population level diet of sea otters represented commercially important prey. Sea otters targeted commercially important species, specifically red sea urchins and Dungeness crab, when first colonizing an area, after which the diet of sea otters became more diverse as colonization durations increased. Using habitat models based on a bivariate normal probability distribution function, environmental covariates and subsistence hunting pressure on sea otters, I determined that sea otter range expansion was limited by subsistence hunting. Further, female and non-territorial males segregated based on habitat and likely prey preferences. I conclude that sea otter populations will likely continue to grow, and that current shellfisheries cannot coexist with sea otters under existing management. Further, conservation and management of sea otter populations, whether to increase the distribution through translocation efforts or reduce the distribution to avoid human conflicts, could benefit from insights gained from spatially explicit modeling at the landscape level.

The impetus for this volume comes from two sources. The first is scientific: by virtue of a preference for certain large benthic invertebrates as food, sea otters have interesting and significant effects on the structure and dynamics of nearshore communities in the North Pacific. The second is political: be cause of the precarious status of the sea otter population in coastal California, the U.S. Fish and Wildlife Service (USFWS) announced, in June 1984, a proposal to establish a new population of sea otters at San Nicolas Island, off southern California. The proposal is based on the premise that risks of catastrophic losses of sea otters, due to large oil spills, are greatly reduced by distributing the population among two geographically separate locations. The federal laws of the U.S. require that USFWS publish an Environmental Impact Statement (ElS) regarding the proposed translocation of sea otters to San Nicolas Island. The EIS is intended to be an assessment of likely bio logical, social, and economic effects of the proposal. In final form, the EIS has an important role in the decision of federal management authority (in this case, the Secretary of the Interior of the U.S.) to accept or reject the proposal.

Sea otter (Enhydra lutris kenyoni) foraging behavior and prey preference (2001-2004) and the behavior and activity budgets of females with dependent pups (2005-2010) were studied during the summer (June-August) in Simpson Bay, Prince William Sound, Alaska. Unlike most previous studies of sea otters which were conducted in coastal areas with a rocky benthos and kelp canopy, the benthic habitat in this study was primarily soft sediment (mud or mixed mud and gravel) with no canopy-forming kelps. Foraging behavior and prey preference. A total of 1,816 foraging dives from 211 bouts were recorded. 87% of foraging dives were successful, and 44% of the prey was identified: 75% clams, 9% Pacific blue mussels, 6% crabs, 2% scallops and a variety of other invertebrates. Significantly more prey items/area were brought up from mixed mud/gravel than mud (p-value

Sea otters and polar bears are carnivorous marine mammals that still resemble their terrestrial ancestors. Compared with Cetacea (whales and dolphins), Sirenia (dugongs and manatees), and Pinnipedia (seals, sea lions, and walrus), they are less adapted for an aquatic life and the most recently evolved among marine mammals. Sea otters are amphibious but seldom come ashore, and polar bears primarily occur on sea ice or along the shore. When at sea, both species spend most of their time swimming at the surface or making short, shallow dives when foraging or pursuing prey. Indeed, polar bears rarely pursue seals in water. Nevertheless, polar bears are powerful swimmers and will stalk seals from the water. As with many other large carnivores, they are solitary hunters. Although sea otters are gregarious and form aggregations at sea called rafts, they are primarily asocial. Except during mating, the principal interaction among sea otters occurs between a female and offspring during the six-month dependency period. In large carnivores (e.g., wolves and lions) that feed on ungulates, sociality and cooperation are favored because of the need to capture large prey and defend carcasses. Polar bears, which are the largest terrestrial carnivore, are solitary hunters of seals and are neither gregarious nor social. Males and females briefly associate during courtship and mating. During this time, males aggressively compete for females. At other times, males generally avoid each other except for aggregations of males that form while summering on land, and females with cubs avoid males, which are known for infanticide. As with sea otters, the interaction of polar bears outside of mating occurs between a female and her offspring during the 2-3 year dependency period. This interaction is critically important when altricial cubs are born in the winter den. This book provides new insight into the ethology and behavioral ecology of sea otters and polar bears. Each chapter reviews the discoveries of previous studies and integrates recent research using new techniques and technology. The authors also address historic and current anthropogenic challenges for their survival as climate change alters entire marine ecosystems.