Consumer and predator foraging behavior can impart profound trait-mediated constraints on community regulation that scale up to influence the structure and stability of ecosystems. Here, we demonstrate how the behavioral response of an apex predator to changes in prey behavior and condition can dramatically alter the role and relative contribution of top-down forcing, depending on the spatial organization of ecosystem states. In 2014, a rapid and dramatic decline in the abundance of a mesopredator (Pycnopodia helianthoides) and primary producer (Macrocystis pyrifera) coincided with a fundamental change in purple sea urchin (Strongylocentrotus purpuratus) foraging behavior and condition, resulting in a spatial mosaic of kelp forests interspersed with patches of sea urchin barrens. We show that this mosaic of adjacent alternative ecosystem states led to an increase in the number of sea otters (Enhydra lutris nereis) specializing on urchin prey, a population-level increase in urchin consumption, and an increase in sea otter survivorship. We further show that the spatial distribution of sea otter foraging efforts for urchin prey was not directly linked to high prey density but rather was predicted by the distribution of energetically profitable prey. Therefore, we infer that spatially explicit sea otter foraging enhances the resistance of remnant forests to overgrazing but does not directly contribute to the resilience (recovery) of forests. These results highlight the role of consumer and predator trait-mediated responses to resource mosaics that are common throughout natural ecosystems and enhance understanding of reciprocal feedbacks between top-down and bottom-up forcing on the regional stability of ecosystems.

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Despite more than a century of federal protection, the California sea otter Enhydra lutris nereis remains threatened under the U.S. Endangered Species Act (ESA), and the population has not appreciably expanded its range in two decades. Here, we examine a novel dataset of 725 sea otter live strandings from 1984-2015 to gain insights into demographic and environmental factors underlying threats to sea otter recovery. Using multinomial logistic regression to evaluate spatiotemporal patterns of stranding causes, we demonstrate that increases in stranding rates, particularly outside the range center, are related to a substantial increase in shark bites. By contrast, trauma linked to human activities has declined dramatically, and now accounts for less than 5% of stranding cases. Within the range core, where the sea otter population seems regulated by prey availability, symptoms of energetic stress represent more than 63% of all strandings and are strongly associated with high sea otter density. Conversely, in range peripheries, the majority of strandings are caused by shark bite and neurological disease. Notably, these threats are virtually absent where nearshore habitat is characterized by at least 10% kelp canopy cover. Our analyses reveal that declining kelp cover may therefore constrain the population's spatial expansion and recovery in two key ways. Absence of kelp intensifies density-independent threats in the range peripheries, and likely limits dispersal of reproductive females, which depend on kelp canopy for nursery habitat. These results highlight the significance of both top-down and bottomup processes in population dynamics, and inform an ecosystem-based approach to conservation planning.

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Translocation and rehabilitation programmes are critical tools for wildlife conservation. These methods achieve greater impact when integrated in a combined strategy for enhancing population or ecosystem restoration. During 2002–2016 we reared 37 orphaned southern sea otter Enhydra lutris nereis pups, using captive sea otters as surrogate mothers, then released them into a degraded coastal estuary. As a keystone species, observed increases in the local sea otter population unsurprisingly brought many ecosystem benefits. The role that surrogate-reared otters played in this success story, however, remained uncertain. To resolve this, we developed an individual-based model of the local population using surveyed individual fates (survival and reproduction) of surrogate-reared and wild-captured otters, and modelled estimates of immigration. Estimates derived from a decade of population monitoring indicated that surrogate-reared and wild sea otters had similar reproductive and survival rates. This was true for males and females, across all ages (1–13 years) and locations evaluated. The model simulations indicated that reconstructed counts of the wild population are best explained by surrogate-reared otters combined with low levels of unassisted immigration. In addition, the model shows that 55% of observed population growth over this period is attributable to surrogate-reared otters and their wild progeny. Together, our results indicate that the integration of surrogacy methods and reintroduction of juvenile sea otters helped establish a biologically successful population and restore a once-impaired ecosystem.