Interactions among sea otters, sea stars, and suspension-feeding invertebrates in the western Aleutian archipelago

The structuring and organizing effects of apex predators on ecosystems are becoming increasingly well documented. The enhancement of kelp forests via sea otter predation on herbivorous sea urchins is among the earliest and best known examples. This study provides evidence for direct and indirect trophic interactions among sea otters, predatory sea stars, and filter-feeding mussels (Mytilus trossulus) and barnacles (Semibalanus cariosis). In western Massacre Bay at Attu Island (173°E, 53°N), subtidal transects showed sea star body size and biomass density declined markedly between 1983 and 1994 as sea otters reinhabited this area. Mussels and barnacles translocated from the rocky intertidal zone to shallow subtidal habitats to assess loss rates from sea star predation showed lower mortality rates after the arrival of sea otters. Prey mortality rates in subtidal caged controls were consistently low and similar to those of intertidal controls in both years. These findings elucidate a trophic pathway by which sea otters can influence ecosystems separate from the well-known sea otter/sea urchin/macroalgae cascade.     

Setting Realistic Recovery Targets for Two Interacting Endangered Species,Sea Otter and Northern Abalone

Failure to account for interactions between endangered species may lead to unexpected population dynamics, inefficient management strategies, waste of scarce resources, and, at worst, increased extinction risk. The importance of species interactions is undisputed, yet recovery targets generally do not account for such interactions. This shortcoming is a consequence of species‐centered legislation, but also of uncertainty surrounding the dynamics of species interactions and the complexity of modeling such interactions. The northern sea otter (Enhydra lutris kenyoni) and one of its preferred prey, northern abalone (Haliotis kamtschatkana), are endangered species for which recovery strategies have been developed without consideration of their strong predator–prey interactions. Using simulation‐based optimization procedures from artificial intelligence, namely reinforcement learning and stochastic dynamic programming, we combined sea otter and northern abalone population models with functional‐response models and examined how different management actions affect population dynamics and the likelihood of achieving recovery targets for each species through time. Recovery targets for these interacting species were difficult to achieve simultaneously in the absence of management. Although sea otters were predicted to recover, achieving abalone recovery targets failed even when threats to abalone such as predation and poaching were reduced. A management strategy entailing a 50% reduction in the poaching of northern abalone was a minimum requirement to reach short‐term recovery goals for northern abalone when sea otters were present. Removing sea otters had a marginally positive effect on the abalone population but only when we assumed a functional response with strong predation pressure. Our optimization method could be applied more generally to any interacting threatened or invasive species for which there are multiple conservation objectives. Definición de Metas de Recuperación Realistas para Dos Especies en Peligro Interactuantes, Enhydra lutris y Haliotis kamtschatkana     

Sea otter foraging on deep-burrowing bivalves in a California coastal lagoon

Sea otter, Enhydra lutris, predation had no detectable effect on abundance and size distribution of deep-burrowing bivalve prey in the Elkhorn Slough, California, USA. Up to 23 otters were present for 6 mo of the study period (March 1984 through April 1985). This is in contrast to previous studies of sea otter predation, especially on the shallow-burrowing Pismo clam Tivela stultorum, which can be found along the wave-exposed coast near the slough. The deep-burrowing clams Tresus nuttallii and Saxidomus nuttalli made up 61% of the prey taken in the slough, and are more difficult for otters to excavate than Pismo clams. The occurrence of foraging otters was highest in an area where the two bivalve prey were extremely abundant (18 individuals m^–2). However, the otters did not selectively prey on the largest clams available within the study sight, but foraged preferentially in a patch of smaller individuals where bivalve burrow depth was restricted by the presence of a dense clay layer. This foraging strategy maximized the amount of prey biomass obtained per unit volume of sediment excavated. Our findings suggest that in soft-sediment habitats deep-burrowing bivalves may be more resistant to otter predation than shallower burrowers.     

Revisiting the classics: considering nonconsumptive effects in textbook examples of predator-prey interactions

Predator effects on prey dynamics are conventionally studied by measuring changes in prey abundance attributed to consumption by predators. We revisit four classic examples of predator-prey systems often cited in textbooks and incorporate subsequent studies of nonconsumptive effects of predators (NCE), defined as changes in prey traits (e.g., behavior, growth, development) measured on an ecological time scale. Our review revealed that NCE were integral to explaining lynx-hare population dynamics in boreal forests, cascading effects of top predators in Wisconsin lakes, and cascading effects of killer whales and sea otters on kelp forests in nearshore marine habitats. The relative roles of consumption and NCE of wolves on moose and consequent indirect effects on plant communities of Isle Royale depended on climate oscillations. Nonconsumptive effects have not been explicitly tested to explain the link between planktonic alewives and the size structure of the zooplankton, nor have they been invoked to attribute keystone predator status in intertidal communities or elsewhere. We argue that both consumption and intimidation contribute to the total effects of keystone predators, and that characteristics of keystone consumers may differ from those of predators having predominantly NCE. Nonconsumptive effects are often considered as an afterthought to explain observations inconsistent with consumption-based theory. Consequently, NCE with the same sign as consumptive effects may be overlooked, even though they can affect the magnitude, rate, or scale of a prey response to predation and can have important management or conservation implications. Nonconsumptive effects may underlie other classic paradigms in ecology, such as delayed density dependence and predator-mediated prey coexistence. Revisiting classic studies enriches our understanding of predator-prey dynamics and provides compelling rationale for ramping up efforts to consider how NCE affect traditional predator-prey models based on consumption, and to compare the relative magnitude of consumptive and NCE of predators.     

Using Ecological Function to Develop Recovery Criteria for Depleted Species: Sea Otters and Kelp Forests in the Aleutian Archipelago

Abstract: Recovery criteria for depleted species or populations normally are based on demographic measures, the goal being to maintain enough individuals over a sufficiently large area to assure a socially tolerable risk of future extinction. Such demographically based recovery criteria may be insufficient to restore the functional roles of strongly interacting species. We explored the idea of developing a recovery criterion for sea otters (Enhydra lutris) in the Aleutian archipelago on the basis of their keystone role in kelp forest ecosystems. We surveyed sea otters and rocky reef habitats at 34 island‐time combinations. The system nearly always existed in either a kelp‐dominated or deforested phase state, which was predictable from sea otter density. We used a resampling analysis of these data to show that the phase state at any particular island can be determined at 95% probability of correct classification with information from as few as six sites. When sea otter population status (and thus the phase state of the kelp forest) was allowed to vary randomly among islands, just 15 islands had to be sampled to estimate the true proportion that were kelp dominated (within 10%) with 90% confidence. We conclude that kelp forest phase state is a more appropriate, sensitive, and cost‐effective measure of sea otter recovery than the more traditional demographically based metrics, and we suggest that similar approaches have broad potential utility in establishing recovery criteria for depleted populations of other functionally important species.     

Serial depletion of marine invertebrates leads to the decline of a strongly interacting grazer.

We investigated the relative roles of natural factors and shoreline harvest leading to recent declines of the black leather chiton (Katharina tunicata) on the outer Kenai Peninsula, Alaska (U.S.A.). This intertidal mollusk is a strongly interacting grazer and a culturally important subsistence fishery for Sugpiaq (Chugach Alutiiq) natives. We took multiple approaches to determine causes of decline. Field surveys examined the significant predictors of Katharina density and biomass across 11 sites varying in harvest pressure, and an integrated analysis of archaeological faunal remains, historical records, traditional ecological knowledge, and contemporary subsistence invertebrate landings examined changes in subsistence practices through time. Strong evidence suggests that current spatial variation in Katharina density and biomass is driven by both human exploitation and sea otter (Enhydra lutris) predation. Traditional knowledge, calibrated by subsistence harvest data, further revealed that several benthic marine invertebrates (sea urchin, crab, clams, and cockles) have declined serially beginning in the 1960s, with reduced densities and sizes of Katharina being the most recent. The timing of these declines was coincident with changes in human behavior (from semi-nomadic to increasingly permanent settlement patterns, improved extractive technologies, regional commercial crustacean exploitation, the erosion of culturally based season and size restrictions) and with the reestablishment of sea otters. We propose that a spatial concentration in shoreline collection pressure through time, increased harvest efficiency, and the serial depletion of alternative marine invertebrate prey have led to intensified per capita predator impacts on Katharina and thus its recent localized decline.     

Effects of river otter activity on terrestrial plants in trophically altered Yellowstone Lake

Animals that deposit aquatically derived nutrients on terrestrial landscapes link food webs and affect a variety of in situ processes. This phenomenon, however, is poorly documented in freshwater habitats, especially where species introductions have drastically changed an ecosystem's trophic structure. In this study, we used stable isotopes to document water-to-land nutrient transport by river otters (Lontra canadensis) around Yellowstone Lake, an ecosystem recently altered by nonnative species invasions. We then investigated the effects of otter fertilization on plant growth and prevalence at latrine (scent-marking) sites and evaluated how the recent changes to the lake's food web could influence these plant responses. Values of delta15N were higher on latrines compared to non-latrine sites in five of seven sample plant taxa. Additionally, latrine grasses had higher percentage N than those from non-latrines. Foliar delta15N positively related to fecal deposition rate for some plants, indicating that increased otter scent-marking led to a rise in these N values. Logistic regression models indicated that otters selected for well-shaded latrines with access to foraging. Atypical latrines, misclassified as non-latrines by the regression models, had values of delta15N similar to correctly classified latrines, suggesting that site effects alone cannot explain elevated N values at otter latrine sites. No difference in plant diversity or percent cover of N-fixing taxa occurred between latrine and nonlatrine sites, though specific genera did differ between site types. Measurements of shoot lengths indicated increased growth of some latrine currants (Ribes sp.). In Yellowstone Lake, a twofold reduction in otter numbers could result in an even greater decline in nutrient deposition at latrines, as otters may become less social in a system with decreased prey availability. Our results highlight the role of animals in linking aquatic and terrestrial habitats in inland freshwater systems and suggest that ongoing changes in the trophic structure of Yellowstone Lake could have unexpected ramifications well beyond the lake itself.     

The influence of size-specific indirect interactions in predator-prey systems

Nonlethal indirect interactions between predators often lead to nonadditive effects of predator number on prey survival and growth. Previous studies have focused on systems with at least two different predator species and one prey species. However, most predators undergo extreme ontological changes in phenotype such that interactions between different-sized cohorts of a predator and its prey could lead to nonadditive effects in systems with only two species. This may be important since different-sized individuals of the same species can differ more in their ecology than similar-sized individuals of different species. This study examined trait-mediated indirect effects in a two-species system including a cannibalistic predator with different-sized cohorts and its prey. I tested for these effects using larvae of two stream salamanders, Gyrinophilus porphyriticus (predator) and Eurycea cirrigera (prey), by altering the densities and combinations of predator size classes in experimental streams. Results showed that the presence of large individuals can significantly reduce the impact of density changes of smaller conspecifics on prey survival through nonlethal means. In the absence of large conspecifics, an increase in the relative frequency of small predators significantly increased predation rates, thereby reducing prey survival. However, with large conspecifics present, increasing the density of small predators did not decrease prey survival, resulting in a 14.3% lower prey mortality than predicted from the independent effects of both predator size classes. Small predators changed their microhabitat use in the presence of larger conspecifics. Prey individuals reduced activity in response to large predators but did not respond to small predators. Both predators reduced prey growth. These results demonstrate that the impact of a predator can be significantly altered by two different types of trait-mediated indirect effects in two-species systems: between different-sized cohorts and between different cohorts and prey. This study demonstrates that predictions based on simple numerical changes that assume independent effects of different size classes or ignore size structure can be strongly misleading. We need to account for the size structure within predator populations in order to predict how changes in predator abundance will affect predator-prey dynamics.     

Indirect effects of prey swamping: differential seed predation during a bamboo masting event

Resource pulses often involve extraordinary increases in prey availability that "swamp" consumers and reverberate through indirect interactions affecting other community members. We developed a model that predicts predator-mediated indirect effects induced by an epidemic prey on co-occurring prey types differing in relative profitability/preference and validated our model by examining current-season and delayed effects of a bamboo mass seeding event on seed survival of canopy tree species in mixed Patagonian forests. The model shows that predator foraging behavior, prey profitability, and the scale of prey swamping influence the character and strength of short-term indirect effects on various alternative prey. When in large prey-swamped patches, nonselective predators decrease predation on all prey types. Selective predators, instead, only benefit prey of similar quality to the swamping species, while very low or high preference prey remain unaffected. Negative indirect effects (apparent competition) may override such positive effects (apparent mutualism), especially for highly preferred prey, when prey-swamped patches are small enough to allow predator aggregation and/or predators show a reproductive numerical response to elevated food supply. Seed predation patterns during bamboo (Chusquea culeou) masting were consistent with predicted short-term indirect effects mediated by a selective predator foraging in large prey-swamped patches. Bamboo seeds and similarly-sized Austrocedrus chilensis (ciprés) and Nothofagus obliqua (roble) seeds suffered lower predation in bamboo flowered than nonflowered patches. Predation rates on the small-seeded Nothofagus dombeyi (coihue) and the large-seeded Nothofagus alpina (rauli) were independent of bamboo flowering. Indirect positive effects were transient; three months after bamboo seeding, granivores preyed heavily upon all seed types, irrespective of patch flowering condition. Moreover, one year after bamboo seeding, predation rates on the most preferred seed (rauli) was higher in flowered than in nonflowered patches. Despite rapid predator numerical responses, short-term positive effects can still influence community recruitment dynamics because surviving seeds may find refuge beneath the litter produced by bamboo dieback. Together, our theoretical analysis and experiments indicate that indirect effects experienced by alternative prey during and after prey-swamping episodes need not be universal but can change across a prey quality spectrum, and they critically depend on predator-foraging rules and the spatial scale of swamping.     

Exploiting sea otter populations: A simulation analysis

We used a Leslie matrix population model to investigate the impact of a range of harvest rates proposed for Alaskan sea otters (Enhydra lutris). The simulation included an analysis of several population mechanisms that might be important in the natural regulation of sea otter populations or in their reactions to harvesting. Significant differences in equilibrium population levels were found between compensatory mechanisms when fixed harvest rates were applied for 25-year periods. Adult harvests set at 2 and 4% of the total population showed that new stable population levels were rapidly attained. Harvest rates of 8 and 10%, however, resulted in marked population declines in simulated harvests. This analysis demonstrates that limited harvesting can be sustained by the population and that otter population compensation responses will be a critical determinant of sustainable harvest rates of sea otter populations.     

Impact of cannibalism on predator-prey dynamics: size-structured interactions and apparent mutualism

Direct and indirect interactions between two prey species can strongly alter the dynamics of predator-prey systems. Most predators are cannibalistic, and as a consequence, even systems with only one predator and one prey include two prey types: conspecifics and heterospecifics. The effects of the complex direct and indirect interactions that emerge in such cannibalistic systems are still poorly understood. This study examined how the indirect interaction between conspecific and heterospecific prey affects cannibalism and predation rates and how the direct interactions between both species indirectly alter the effect of the cannibalistic predator. I tested for these effects using larvae of the stream salamanders Eurycea cirrigera (prey) and Pseudotriton ruber (cannibalistic predator) by manipulating the relative densities of the conspecific and heterospecific prey in the presence and absence of the predator in experimental streams. The rates of cannibalism and heterospecific predation were proportional to the respective densities and negatively correlated, indicating a positive indirect interaction between conspecific and heterospecific prey, similar to "apparent mutualism." Direct interactions between prey species did not alter the effect of the predator. Although both types of prey showed a similar 30% reduction in night activity and switch in microhabitat use in response to the presence of the predator, cannibalism rates were three times higher than heterospecific predation rates irrespective of the relative densities of the two types of prey. Cumulative predation risks differed even more due to the 48% lower growth rate of conspecific prey. Detailed laboratory experiments suggest that the 3:1 difference in cannibalism and predation rate was due to the higher efficiency of heterospecific prey in escaping immediate attacks. However, no difference was observed when the predator was a closely related salamander species, Gyrinophilus porphyriticus, indicating that this difference is species specific. This demonstrates that cannibalism can result in the coupling of predator and prey mortality rates that strongly determines the dynamics of predator-prey systems.     

Can rare positive interactions become common when large carnivores consume livestock?

Livestock populations in protected areas are viewed negatively because of their interaction with native ungulates through direct competition for food resources. However, livestock and native prey can also interact indirectly through their shared predator. Indirect interactions between two prey species occur when one prey modifies either the functional or numerical responses of a shared predator. This interaction is often manifested as negative effects (apparent competition) on one or both prey species through increased predation risk. But indirect interactions can also yield positive effects on a focal prey if the shared predator modifies its functional response toward increased consumption of an abundant and higher-quality alternative prey. Such a phenomenon between two prey species is underappreciated and overlooked in nature. Positive indirect effects can be expected to occur in livestock-dominated wildlife reserves containing large carnivores. We searched for such positive effects in Acacia-Zizhypus forests of India's Gir sanctuary where livestock (Bubalus bubalis and Bos indicus) and a coexisting native prey (chital deer, Axis axis) are consumed by Asiatic lions (Panthera leo persica). Chital vigilance was higher in areas with low livestock density than in areas with high livestock density. This positive indirect effect occurred because lion predation rates on livestock were twice as great where livestock were abundant than where livestock density was low. Positive indirect interactions mediated by shared predators may be more common than generally thought with rather major consequences for ecological understanding and conservation. We encourage further studies to understand outcomes of indirect interactions on long-term predator-prey dynamics in livestock-dominated protected areas.     

Constancy and change in marine predator diets across a shift in oceanographic conditions in the Northern California Current

Variable ocean conditions can greatly impact prey assemblages and predator foraging in marine ecosystems. Our goal was to better understand how a change in ocean conditions influenced dietary niche overlap among a suite of midtrophic-level predators. We examined the diets of three fishes and one seabird off central Oregon during two boreal summer upwelling periods with contrasting El Niño (2010) and La Niña (2011) conditions. We found greater niche specialization during El Niño and increased niche overlap during La Niña in both the nekton and micronekton diet components, especially in the larger, more offshore predators. However, only the two smaller, more nearshore predators exhibited interannual variation in diet composition. Concurrent trawl surveys confirmed that changes in components of predator diets reflected changes in the prey community. Using multiple predators across diverse taxa and life histories provided a comprehensive understanding of food-web dynamics during changing ocean conditions.     

Consequences of stage-structured predators: cannibalism, behavioral effects, and trophic cascades

Cannibalistic and asymmetrical behavioral interactions between stages are common within stage-structured predator populations. Such direct interactions between predator stages can result in density- and trait-mediated indirect interactions between a predator and its prey. A set of structured predator-prey models is used to explore how such indirect interactions affect the dynamics and structure of communities. Analyses of the separate and combined effects of stage-structured cannibalism and behavior-mediated avoidance of cannibals under different ecological scenarios show that both cannibalism and behavioral avoidance of cannibalism can result in short- and long-term positive indirect connections between predator stages and the prey, including "apparent mutualism." These positive interactions alter the strength of trophic cascades such that the system's dynamics are determined by the interaction between bottom-up and top-down effects. Contrary to the expectation of simpler models, enrichment increases both predator and prey abundance in systems with cannibalism or behavioral avoidance of cannibalism. The effect of behavioral avoidance of cannibalism, however, depends on how strongly it affects the maturation rate of the predator. Behavioral interactions between predator stages reduce the short-term positive effect of cannibalism on the prey density, but can enhance its positive long-term effects. Both interaction types reduce the destabilizing effect of enrichment. These results suggest that inconsistencies between data and simple models can be resolved by accounting for stage-structured interactions within and among species.     

Functional responses: a question of alternative prey and predator density

Throughout the study of ecology, there has been a growing realization that indirect effects among species cause complexity in food webs. Understanding and predicting the behavior of ecosystems consequently depends on our ability to identify indirect effects and their mechanisms. The present study experimentally investigates indirect interactions arising between two prey species that share a common predator. In a natural field experiment, we introduced different densities of mealworms (Tenebrio molitor), an alternative prey, to a previously studied predator-prey system in which paper wasps (Polistes dominulus) preyed on shield beetle larvae (Cassida rubiginosa). We tested if alternative prey affects predation on the first prey (i.e., the predator-dependent functional response of paper wasps) by modifying either interference among predators or the effective number of predators foraging on shield beetles. Presence of mealworms significantly reduced the effective number of predators, whereas predator interference was not affected. In this way, the experimentally introduced alternative prey altered the wasps' functional response and thereby indirectly influenced C. rubiginosa density. In all prey-density combinations offered, paper wasps constantly preferred T. molitor. This led to an asymmetrical, indirect interaction between both prey species: an increase in mealworm density significantly relaxed predation on C. rubiginosa, whereas an increase in C. rubiginosa density intensified predation on mealworms. Such asymmetrical outcomes of a fixed food preference can significantly affect the population dynamics of the species involved. In spite of the repeated finding of a Type III functional response in this system, our experiment did not reveal switching behavior in paper wasps. The variety of mechanisms underlying direct and indirect interactions within our study system exemplifies the importance of incorporating alternative prey when investigating the impact of a generalist predator on a focal prey population under realistic field conditions.     

Lethal and nonlethal predator effects on an herbivore guild mediated by system productivity

Indirect effects propagated through intervening species in a food web have important effects on community properties. Traditionally, these indirect effects have been conceptualized as mediated through density changes of the intervening species, but it is becoming increasingly apparent that those mediated through trait (phenotypic) responses also can be very important. Because density- and trait-mediated indirect effects have different properties, it is critical that we understand the mechanisms of transmission in order to predict how they will interact, and when or where they will be important. In this study, we examined the mechanisms and consequences of the lethal (density-mediated) and nonlethal (trait-mediated) effects of a larval odonate predator on a guild of four herbivore species (a larval anuran and three species of snails) and their resources. We also manipulated system productivity in order to explore the effects of environmental context on the transmission of these two types of indirect effects. We show that trait-mediated effects arising from the predator can be very strong relative to density-mediated effects on both the competing herbivores and the species composition and production of their resources. A number of these indirect effects are shown to be contingent on productivity of the system. We further present evidence that trait- and density-mediated indirect effects originating from a predator may be transmitted independently through different routes in a food web, particularly when spatial responses of the transmitting prey are involved. Finally, effects on prey growth due to trait responses to the predator varied from negative to positive in predictable ways as a function of time and indirect effects on the larger food web. These results indicate the important role that trait-mediated indirect effects can play in trophic cascades and keystone predator interactions, and we discuss how the mechanisms involved can be incorporated in theory.     

Conservation Strategies for Species Affected by Apparent Competition

Apparent competition is an indirect interaction between 2 or more prey species through a shared predator, and it is increasingly recognized as a mechanism of the decline and extinction of many species. Through case studies, we evaluated the effectiveness of 4 management strategies for species affected by apparent competition: predator control, reduction in the abundances of alternate prey, simultaneous control of predators and alternate prey, and no active management of predators or alternate prey. Solely reducing predator abundances rapidly increased abundances of alternate and rare prey, but observed increases are likely short‐lived due to fast increases in predator abundance following the cessation of control efforts. Substantial reductions of an abundant alternate prey resulted in increased predation on endangered huemul (Hippocamelus bisulcus) deer in Chilean Patagonia, which highlights potential risks associated with solely reducing alternate prey species. Simultaneous removal of predators and alternate prey increased survival of island foxes (Urocyon littoralis) in California (U.S.A.) above a threshold required for population recovery. In the absence of active management, populations of rare woodland caribou (Rangifer tarandus caribou) continued to decline in British Columbia, Canada. On the basis of the cases we examined, we suggest the simultaneous control of predators and alternate prey is the management strategy most likely to increase abundances and probabilities of persistence of rare prey over the long term. Knowing the mechanisms driving changes in species’ abundances before implementing any management intervention is critical. We suggest scientists can best contribute to the conservation of species affected by apparent competition by clearly communicating the biological and demographic forces at play to policy makers responsible for the implementation of proposed management actions. Estrategias de Conservación para Especies Afectadas por Competencia Aparente     

Differential tree and shrub production in response to fertilization and disturbance by coastal river otters in Alaska

We explored the interacting effects of marine-derived nutrient fertilization and physical disturbance introduced by coastal river otters (Lontra canadensis) on the production and nutrient status of pristine shrub and tree communities in Prince William Sound, Alaska, USA. We compared production of trees and shrubs between latrines and non-latrines, while accounting for otter site selection, by sampling areas on and off sites. Nitrogen stable isotope analysis (delta15N) indicated that dominant tree and shrub species assimilated the marine-derived N excreted by otters. In association with this uptake, tree production increased, but shrub density and nonwoody aboveground shrub production decreased. The reduced shrub production was caused by destruction of ramets, especially blueberry (Vaccinium spp.), through physical disturbance by river otters. False azalea (Menziesia ferruginea) ramets were less sensitive to otter disturbance. Although surviving individual blueberry ramets showed a tendency for increased production per plant, false azalea allocated excess N to storage in leaves rather than growth. We found that plant responses to animal activity vary among species and levels of biological organization (leaf, plant, ecosystem). Such differences should be accounted for when assessing the influence of river otters on the carbon budget of Alaskan coastal forests at the landscape scale.     

Incorporating diverse data and realistic complexity into demographic estimation procedures for sea otters.

Reliable information on historical and current population dynamics is central to understanding patterns of growth and decline in animal populations. We developed a maximum likelihood-based analysis to estimate spatial and temporal trends in age/sex-specific survival rates for the threatened southern sea otter (Enhydra lutris nereis), using annual population censuses and the age structure of salvaged carcass collections. We evaluated a wide range of possible spatial and temporal effects and used model averaging to incorporate model uncertainty into the resulting estimates of key vital rates and their variances. We compared these results to current demographic parameters estimated in a telemetry-based study conducted between 2001 and 2004. These results show that survival has decreased substantially from the early 1990s to the present and is generally lowest in the north-central portion of the population's range. The greatest temporal decrease in survival was for adult females, and variation in the survival of this age/sex class is primarily responsible for regulating population growth and driving population trends. Our results can be used to focus future research on southern sea otters by highlighting the life history stages and mortality factors most relevant to conservation. More broadly, we have illustrated how the powerful and relatively straightforward tools of information-theoretic-based model fitting can be used to sort through and parameterize quite complex demographic modeling frameworks.     

Prey defense,predator preference,and nonrandom diet: The interactions between Pycnopodia helianthoides and two species of sea urchins

Interactions between the predatory sea star Pycnopodia helianthoides (Brandt, 1835) and two of its natural prey, the sea urchins Strongylocentrotus purpuratus (Stimpson, 1857) and S. franciscanus (Agassiz, 1863), are examined with regard to predator preference, predator diet, and prey defenses. The sea star is able to detect both species of sea urchin upstream in a Y-trough, but does not consistently choose one over the other (i.e., no preference). However, when the sea star is presented with equal numbers of similar-sized specimens of the two species of sea urchin, its diet is markedly nonrandom, since S. purpuratus is eaten almost 98% of the time. The defensive responses of the two species of sea urchin differ in form and effectiveness. S. franciscanus employs its long spines as defensive weapons, pinching the rays of an attacking sea star. This defensive response is more effective than the pedicellarial response used by S. purpuratus. The nonrandom diet of the predator seems to result primarily from prey defensive responses that differ in effectiveness, rather than from an intrinsic, behavioral preference of the predator at an earlier stage in the predator/prey interaction.