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J. A. Estes

Publications and source records attributed to J. A. Estes.

At least 19 recordsLinked to original sources

Changes in abiotic drivers of green sea urchin demographics following the loss of a keystone predator

Sea urchin population demographics can respond to changes in keystone species abundances, with the magnitude of these responses varying depending on environmental influences. In this study, sea urchin populations were surveyed across 15 Aleutian archipelago islands over a 30-year period to understand how patterns of sea urchin demography (density, biomass, and size structure) varied through different ecological regimes that were caused by changes in the abundance of sea otters, a keystone species in this system. To examine long-term changes in sea urchin demographics, four time periods across the recent decline of sea otters were examined: during sea otter presence (1987-1994), nearing absence at the end of the decline (1997-2000), 10 years postdecline (2008-2010), and 15-20 years following the loss of sea otters from the ecosystem (2014-2017). Our results show that when sea otters were broadly present, sea urchin demographics were generally similar across the archipelago, with few urchins that had large-sized bodies. During this time, bottom-up environmental controls were muted relative to top-down forces from keystone predation. However, as sea otters declined and remained absent from the system, abiotic factors became more influential on sea urchin biomass, density, and size structure. In particular, differences among island groups during these periods were correlated with variation in ocean temperature, bathymetric complexity, and habitat availability. Sea urchin recruitment also varied among island groups, corresponding to ecoregions delineated by oceanic passes across the archipelago. The functional extinction of sea otters revealed an increasing influence of abiotic forcing in the absence of top-down control. This study further highlights the importance of understanding how keystone predators regulate herbivore demographics.

Alaska

Ontogenetic and among-individual variation in foraging strategies of northeast Pacific white sharks based on stable isotope analysis

There is growing evidence for individuality in dietary preferences and foraging behaviors within populations of various species. This is especially important for apex predators, since they can potentially have wide dietary niches and a large impact on trophic dynamics within ecosystems. We evaluate the diet of an apex predator, the white shark ( Carcharodon carcharias ), by measuring the stable carbon and nitrogen isotope composition of vertebral growth bands to create lifetime records for 15 individuals from California. Isotopic variations in white shark diets can reflect within-region differences among prey (most importantly related to trophic level), as well as differences in baseline values among the regions in which sharks forage, and both prey and habitat preferences may shift with age. The magnitude of isotopic variation among sharks in our study (>5‰ for both elements) is too great to be explained solely by geographic differences, and so must reflect differences in prey choice that may vary with sex, size, age and location. Ontogenetic patterns in δ 15 N values vary considerably among individuals, and one third of the population fit each of these descriptions: 1) δ 15 N values increased throughout life, 2) δ 15 N values increased to a plateau at ~5 years of age, and 3) δ 15 N values remained roughly constant values throughout life. Isotopic data for the population span more than one trophic level, and we offer a qualitative evaluation of diet using shark-specific collagen discrimination factors estimated from a 3+ year captive feeding experiment (Δ 13 C shark-diet and Δ 15 N shark-diet equal 4.2‰ and 2.5‰, respectively). We assess the degree of individuality with a proportional similarity index that distinguishes specialists and generalists. The isotopic variance is partitioned among differences between-individual (48%), within-individuals (40%), and by calendar year of sub-adulthood (12%). Our data reveal substantial ontogenetic and individual dietary variation within a white shark population.

PLoS ONE

Predicting community responses to perturbations in the face of imperfect knowledge and network complexity

How best to predict the effects of perturbations to ecological communities has been a long-standing goal for both applied and basic ecology. This quest has recently been revived by new empirical data, new analysis methods, and increased computing speed, with the promise that ecologically important insights may be obtainable from a limited knowledge of community interactions. We use empirically based and simulated networks of varying size and connectance to assess two limitations to predicting perturbation responses in multispecies communities: (1) the inaccuracy by which species interaction strengths are empirically quantified and (2) the indeterminacy of species responses due to indirect effects associated with network size and structure. We find that even modest levels of species richness and connectance (∼25 pairwise interactions) impose high requirements for interaction strength estimates because system indeterminacy rapidly overwhelms predictive insights. Nevertheless, even poorly estimated interaction strengths provide greater average predictive certainty than an approach that uses only the sign of each interaction. Our simulations provide guidance in dealing with the trade-offs involved in maximizing the utility of network approaches for predicting dynamics in multispecies communities.

Ecology

Sea otter mortality in fish and shellfish traps: Estimating potential impacts and exploring possible solutions

Sea otters Enhydra lutris can be bycaught and drowned in fishing pots and traps, which may pose a threat to the welfare of otter populations. We explored this potential problem and its solutions using a wide variety of analyses. We exposed live California (USA) sea otters to finfish traps, lobster traps, and mock Dungeness crab traps in captive trials and found that the animals attempted to enter the circular and rectangular fyke openings, with some becoming entrapped. Using both live and dead sea otters, we found that a 3 × 9 inch (7.6 × 22.9 cm) fyke opening (1 inch narrower than the 4 × 9 inch [10.2 × 22.9 cm] openings currently used in California’s commercial Dungeness crab fishery) would exclude most free-living (i.e. weaned from their mothers) otters while permitting the undiminished capture of crabs. Observer programs do not currently exist in California for these fisheries, so we calculated the effort required by an observer program to document sea otter bycatch over a range of hypothetical levels and evaluated the impact of those mortality rates on population growth. These analyses demonstrate that significant mortality from bycatch might easily go undetected, even with seemingly high levels of observer effort. As sea otters reoccupy portions of their former habitat in California, co-occurrence with finfish and shellfish traps with relatively large fyke openings will increase.

Endangered Species Research

Using stable isotopes to investigate individual diet specialization in California sea otters (Enhydra lutris nereis)

Differences in diet composition among conspecifics (dietary specialization) have been documented across a broad range of taxonomic groups and habitats, and such variation at the individual level is increasingly recognized as an important component of diversity in trophic interactions. Accurate identification of individual dietary specialization, however, requires longitudinal dietary records that are labor-intensive and cost-prohibitive to obtain for many species. Here we explore the use of stable isotopes (δ 13 C and δ 15 N) as a promising technique for detecting and quantifying patterns of individual dietary specialization. Southern sea otters ( Enhydra lutris nereis ) offer a unique opportunity for testing this approach because (1) they consume a wide variety of prey that span multiple trophic levels, habitats, and ecologically defined functional groups; and (2) individual diet specialization can be validated with existing observational data. We analyzed the isotopic composition of sea otter vibrissae ( n = 31) in order to characterize inter- and intra-individual variation in sea otter diets at Monterey Bay, California, USA. At the population level, sea otters showed substantial variation in both δ 13 C and δ 15 N values, occupying nearly all of the “isotopic space” created by the diversity of isotopic signatures of potential prey taxa. Most of the variation in sea otter vibrissae was accounted for by differences between individuals, with much less contributed by within-individual variation. A majority of sea otters (∼80%) showed relatively little temporal variability in isotopic composition, suggesting that the proportional composition of most individuals' diets is relatively constant over time; a few individuals (∼20%) exhibited a high degree of intra-vibrissa isotopic variability, suggesting seasonal shifts in diet composition. These results and our interpretation of them were supported by long-term observational data on the diets of radio-tagged sea otters from the same population ( n = 23). Our results demonstrate that stable isotopes can provide an efficient tool for measuring individual- and population-level dietary breadth and may be useful for studying populations where longitudinal data on individuals would otherwise be impossible to acquire. This will be critical for examining the causes and consequences of dietary variation within and among consumer populations, thereby improving our understanding of these important ecological and evolutionary processes at the community level.

Ecology

Learning to be different: Acquired skills, social learning, frequency dependence, and environmental variation can cause behaviourally mediated foraging specializations

Question: How does the ability to improve foraging skills by learning, and to transfer that learned knowledge, affect the development of intra-population foraging specializations? Features of the model: We use both a state-dependent life-history model implemented by stochastic dynamic programming (SDPM) and an individual-based model (IBM) to capture the dynamic nature of behavioural preferences in feeding. Variables in the SDPM include energy reserves, skill levels, energy and handling time per single prey item, metabolic rate, the rates at which skills are learned and forgotten, the effect of skills on handling time, and the relationship between energy reserves and fitness. Additional variables in the IBM include the probability of successful weaning, the logistic dynamics of the prey species with stochastic recruitment, the intensity of top-down control of prey by predators, the mean and variance in skill levels of new recruits, and the extent to which learned Information can be transmitted via matrilineal social learning. Key range of variables: We explore the effects of approaching the time horizon in the SDPM, changing the extent to which skills can improve with experience, increasing the rates of learning or forgetting of skills, changing whether the learning curve is constant, accelerating (T-shaped) or decelerating ('r'-shaped), changing both mean and maximum possible energy reserves, changing metabolic costs of foraging, and changing the rate of encounter with prey. Conclusions: The model results show that the following factors increase the degree of prey specialization observed in a predator population: (1) Experience handling a prey type can substantially improve foraging skills for that prey. (2) There is limited ability to retain complex learned skills for multiple prey types. (3) The learning curve for acquiring new foraging skills is accelerating, or J-shaped. (4) The metabolic costs of foraging are high relative to available energy reserves. (5) Offspring can learn foraging skills from their mothers (matrilineal social learning). (6) Food abundance is limited, such that average individual energy reserves are low Additionally, the following factors increase the likelihood of alternative specializations co-occurring in a predator population: (1) The predator exerts effective top-down control of prey abundance, resulting in frequency-dependent dynamics. (2) There is stochastic Variation in prey population dynamics, but this Variation is neither too extreme in magnitude nor too 'slow' with respect to the time required for an individual forager to learn new foraging skills. For a given predator population, we deduce that the degree of specialization will be highest for those prey types requiring complex capture or handling skills, while prey species that are both profitable and easy to capture and handle will be included in the diet of all individuals. Frequency-dependent benefits of selecting alternative prey types, combined with the ability of foragers to improve their foraging skills by learning, and transmit learned skills to offspring, can result in behaviourally mediated foraging specialization, and also lead to the co-existence of alternative specializations. The extent of such specialization is predicted to be a variable trait, increasing in locations or years when intra-specific competition is high relative to inter-specific competition. ?? 2009 M. Tim Tinker.

Evolutionary Ecology Research

Bald eagles and sea otters in the Aleutian Archipelago: indirect effects of trophic cascades.

Because sea otters ( Enhydra lutris ) exert a wide array of direct and indirect effects on coastal marine ecosystems throughout their geographic range, we investigated the potential influence of sea otters on the ecology of Bald Eagles ( Haliaeetus leucocephalus ) in the Aleutian Islands, Alaska, USA. We studied the diets, productivity, and density of breeding Bald Eagles on four islands during 1993–1994 and 2000–2002, when sea otters were abundant and scarce, respectively. Bald Eagles depend on nearshore marine communities for most of their prey in this ecosystem, so we predicted that the recent decline in otter populations would have an indirect negative effect on diets and demography of Bald Eagles. Contrary to our predictions, we found no effects on density of breeding pairs on four islands from 1993–1994 to 2000–2002. In contrast, diets and diet diversity of Bald Eagles changed considerably between the two time periods, likely reflecting a change in prey availability resulting from the increase and subsequent decline in sea otter populations. The frequency of sea otter pups, rock greenling ( Hexagammus lagocephalus ), and smooth lumpsuckers ( Aptocyclus ventricosus ) in the eagle's diet declined with corresponding increases in Rock Ptarmigan ( Lagopus mutus ), Glaucous-winged Gulls ( Larus glaucescens ), Atka mackerel ( Pleurogrammus monopterygius ), and various species of seabirds during the period of the recent otter population decline. Breeding success and productivity of Bald Eagles also increased during this time period, which may be due to the higher nutritional quality of avian prey consumed in later years. Our results provide further evidence of the wide-ranging indirect effects of sea otter predation on nearshore marine communities and another apex predator, the Bald Eagle. Although the indirect effects of sea otters are widely known, this example is unique because the food-web pathway transcended five species and several trophic levels in linking one apex predator to another.

Ecology

Environmental contaminants in bald eagle eggs from the Aleutian archipelago

We collected 136 fresh and unhatched eggs from bald eagle (Haliaeetus leucocephalus) nests and assessed productivity on eight islands in the Aleutian archipelago, 2000 to 2002. Egg contents were analyzed for a broad spectrum of organochlorine (OC) contaminants, mercury (Hg), and stable isotopes of carbon (??13C) and nitrogen (??15N). Concentrations of polychlorinated biphenyls (??PCBs), p,p???- dichlorodiphenyldichloroethylene (DDE), and Hg in bald eagle eggs were elevated throughout the archipelago, but the patterns of distribution differed among the various contaminants. Total PCBs were highest in areas of past military activities on Adak and Amchitka Islands, indicating local point sources of these compounds. Concentrations of DDE and Hg were higher on Amchitka Island, which was subjected to much military activity during World War II and the middle of the 20th century. Concentrations of ??PCBs also were elevated on islands with little history of military activity (e.g., Amlia, Tanaga, Buldir), suggesting non-point sources of PCBs in addition to point sources. Concentrations of DDE and Hg were highest in eagle eggs from the most western Aleutian Islands (e.g., Buldir, Kiska) and decreased eastward along the Aleutian chain. This east-to-west increase suggested a Eurasian source of contamination, possibly through global transport and atmospheric distillation and/or from migratory seabirds. Eggshell thickness and productivity of bald eagles were normal and indicative of healthy populations because concentrations of most contaminants were below threshold levels for effects on reproduction. Contrary to our predictions, contaminant concentrations were not correlated with stable isotopes of carbon (??13C) or nitrogen (??15N) in eggs. These latter findings indicate that contaminant concentrations were influenced more by point sources and geographic location than trophic status of eagles among the different islands. ?? 2007 SETAC.

Environmental Toxicology and Chemistry

Individual dietary specialization and dive behaviour in the California sea otter: Using archival time-depth data to detect alternative foraging strategies

The existence of individual prey specializations has been reported for an ever-growing number of taxa, and has important ramifications for our understanding of predator-prey dynamics. We use the California sea otter population as a case study to validate the use of archival time-depth data to detect and measure differences in foraging behaviour and diet. We collected observational foraging data from radio-tagged sea otters that had been equipped with Mk9 time depth recorders (TDRs, Wildlife Computers, Redmond, WA). After recapturing the study animals and retrieving the TDRs it was possible to compare the two data types, by matching individual dives from the TDR record with observational data and thus examining behavioural correlates of capture success and prey species. Individuals varied with respect to prey selection, aggregating into one of three distinct dietary specializations. A number of TDR-derived parameters, particularly dive depth and post-dive surface interval, differed predictably between specialist types. A combination of six dive parameters was particularly useful for discriminating between specialist types, and when incorporated into a multivariate cluster analysis, these six parameters resulted in classification of 13 adult female sea otters into three clusters that corresponded almost perfectly to the diet-based classification (1 out of 13 animals was misclassified). Thus based solely on quantifiable traits of time-depth data that have been collected over an appropriate period (in this case 1 year per animal), it was possible to assign female sea otters to diet type with >90% accuracy. TDR data can thus be used as a tool to measure the degree of individual specialization in sea otter populations, a conclusion that will likely apply to other diving marine vertebrates as well. Our ultimate goals must be both to understand the causes of individual specialization, and to incorporate such variation into models of population- and community-level food web dynamics. ?? 2007 Elsevier Ltd. All rights reserved.

Deep-Sea Research Part II: Topical Studies in Ocea

Causes of mortality in California sea otters during periods of population growth and decline

Elevated mortality appears to be the main reason for both sluggish growth and periods of decline in the threatened California sea otter population . We assessed causes of mortality from salvage records of 3,105 beach-cast carcasses recovered from 1968 through 1999, contrasting two periods of growth with two periods of decline . Overall, an estimated 40%-60% of the deaths were not recovered and 70% of the recovered carcasses died from unknown causes . Nonetheless, several common patterns were evident in the salvage records during the periods of population decline . These included greater percentages of (1) prime age animals (3-10 yr), (2) carcasses killed by great white shark attacks, (3) carcasses recovered in spring and summer, and (4) carcasses for which the cause of death was unknown. Neither sex composition nor the proportion of carcasses dying of infectious disease varied consistently between periods of population increase and decline . The population decline from 1976 to 1984 was likely due to incidental mortality in a set-net fishery, and the decline from 1995 to 1999 may be related to a developing live-fish fishery. Long-term trends unrelated to periods of growth and decline included a decrease in per capita pup production and mass/length ratios of adult carcasses over the 31-yr study. The generally high proportion of deaths from infectious disease suggests that this factor has contributed to the chronically sluggish growth rate of the California sea otter population .

California

Patterns of growth and body condition in sea otters from the Aleutian archipelago before and after the recent population decline

1. Growth models for body mass and length were fitted to data collected from 1842 sea otters Enhydra lutris shot or live-captured throughout south-west Alaska between 1967 and 2004. Growth curves were constructed for each of two main year groups: 1967–71 when the population was at or near carrying capacity and 1992–97 when the population was in steep decline. Analyses of data collected from animals caught during 2004, when the population density was very low, were precluded by a small sample size and consequently only examined incidentally to the main growth curves. 2. Growth curves demonstrated a significant increase in body mass and body length at age in the 1990s. Asymptotic values of body mass were 12–18% higher in the 1990s than in the 1960s/70s, and asymptotic values for body length were 10–11% higher between the same periods. Data collected in 2004 suggest a continued increase in body size, with nearly all data points for mass and length falling significantly above the 1990s growth curves. 3. In addition to larger asymptotic values for mass and length, the rate of growth towards asymptotic values was more rapid in the 1990s than in the 1960s/70s: sea otters reached 95% of asymptotic body mass and body length 1–2 years earlier in the 1990s. 4. Body condition (as measured by the log mass/log length ratio) was significantly greater in males than in females. There was also an increasing trend from the 1960s/70s through 2004 despite much year-to-year variation. 5. Population age structures differed significantly between the 1960s/70s and the 1990s with the latter distribution skewed toward younger age classes (indicating an altered l x function) suggesting almost complete relaxation of age-dependent mortality patterns (i.e. those typical of food-limited populations). 6. This study spanned a period of time over which the population status of sea otters in the Aleutian archipelago declined precipitously from levels at or near equilibrium densities at some islands in the 1960s/70s to < 5% of estimated carrying capacity by the late 1990s. The results of this study indicate an improved overall health of sea otters over the period of decline and suggest that limited nutritional resources were not the cause of the observed reduced population abundance. Our findings are consistent with the hypothesis that the decline was caused by increased killer whale predation.

Journal of Animal Ecology

An introduced predator alters Aleutian Island plant communities by thwarting nutrient subsidies

The ramifying effects of top predators on food webs traditionally have been studied within the framework of trophic cascades. Trophic cascades are compelling because they embody powerful indirect effects of predators on primary production. Although less studied, indirect effects of predators may occur via routes that are not exclusively trophic. We quantified how the introduction of foxes onto the Aleutian Islands transformed plant communities by reducing abundant seabird populations, thereby disrupting nutrient subsidies vectored by seabirds from sea to land. We compared soil and plant fertility, plant biomass and community composition, and stable isotopes of nitrogen in soil, plants, and other organisms on nine fox-infested and nine historically fox-free islands across the Aleutians. Additionally, we experimentally augmented nutrients on a fox-infested island to test whether differences in plant productivity and composition between fox-infested and fox-free islands could have arisen from differences in nutrient inputs between island types. Islands with historical fox infestations had soils low in phosphorus and nitrogen and plants low in tissue nitrogen. Soils, plants, slugs, flies, spiders, and bird droppings on these islands had low d15N values indicating that these organisms obtained nitrogen from internally derived sources. In contrast, soils, plants, and higher trophic level organisms on fox-free islands had elevated d15N signatures indicating that they utilized nutrients derived from the marine environment. Furthermore, soil phosphorus (but not nitrogen) and plant tissue nitrogen were higher on fox-free than fox-infested islands. Nutrient subsidized fox-free islands supported lush, high biomass plant communities dominated by graminoids. Fox-infested islands were less graminoid dominated and had higher cover and biomass of low-lying forbs and dwarf shrubs. While d15N profiles of soils and plants and graminoid biomass varied with island size and distance from shore, after accounting for these effects differences between fox-infested and fox-free islands still existed. Fertilization over four years caused a 24-fold increase in graminoid biomass and a shift toward a more graminoid dominated plant community typical of fox-free islands. These results indicate that apex predators can influence plant productivity and composition through complex interaction web pathways involving both top-down forcing and bottom-up nutrient exchanges across systems. ?? 2006 by the Ecological Society of America.

Ecological Monographs

The ecology of extinctions in kelp forest communities

We recognize three levels of extinction–global, local, and ecological – and provide examples of each. The protection and recovery of sea otters (Enhydra lutris) has provided abundant evidence of the consequences of their local extinction from kelp forest communities in the North Pacifc Ocean. These consequences include release of benthic invertebrate populations from limitation by predation; deforestation of kelp beds due to increased grazing by herbivorous sea urchins, one of the otter's main prey; and various cascading effects resulting from the biological and physical importance of kelp in coastal ecosystems. These interactions probably were important agents of selection for certain species. Two other examples are discussed: Steller's sea cow (Hydrodamalis gigas), a case of global extinction, and spiny lobsters, a possible case of ecological extinction. We speculate that grazing by sea cows was an important disturbance to surface-canopy-forming kelps and other algae in the littoral zones, but also point out that any such interactions probably acted in concert with physical disturbances by ocean waves. The ecological and evolutionary importance of sea cow grazing probably will remain a matter of speculation and conjecture because the species is globally extinct. Predation by spiny lobsters limits a variety of littoral and sublittoral invertebrate populations, particularly mollusks In one remarkable example, the reduction or local extinction of spiny lobsters enabled predutory whelks to increase in size and abundance, ultimately resulting in a predutor-prey role reversal. From these and other case studies we can clued that (1) the extinction of consumers may have brad and sometimes unexpected influences on kelp forest ecosystems; (2) direct or indirect interactions with now-extinct species probably exerted important selective influences on many extant forms; (3) such ecological and evolutionary influences are best understood where local or ecological extinctions, followed by recoveries, have provided comparisons in space or time; and (4) because of various ecological and behavioral barriers, local extinctions and their ecological consequences may not be simply reversed by protecting or reintroducing depleted or locally extinct species.

Conservation Biology

Evolution of large body size in abalones (Haliotis): Patterns and implications

Kelps and other fleshy macroalgae - dominant reef-inhabiting organisms in cool - seasmay have radiated extensively following late Cenozoic polar cooling, thus triggering a chain of evolutionary change in the trophic ecology of nearshore temperate ecosystems. We explore this hypothesis through an analysis of body size in the abalones (Gastropoda; Haliotidae), a widely distributed group in modern oceans that displays a broad range of body sizes and contains fossil representatives from the late Cretaceous (60-75 Ma). Geographic analysis of maximum shell length in living abalones showed that small-bodied species, while most common in the Tropics, have a cosmopolitan distribution, whereas large-bodied species occur exclusively in cold-water ecosystems dominated by kelps and other macroalgae. The phylogeography of body size evolution in extant abalones was assessed by constructing a molecular phylogeny in a mix of large and small species obtained from different regions of the world. This analysis demonstrates that small body size is the plesiomorphic state and largeness has likely arisen at least twice. Finally, we compiled data on shell length from the fossil record to determine how (slowly or suddenly) and when large body size arose in the abalones. These data indicate that large body size appears suddenly at the Miocene/Pliocene boundary. Our findings support the view that fleshy-algal dominated ecosystems radiated rapidly in the coastal oceans with the onset of the most recent glacial age. We conclude with a discussion of the broader implications of this change. ?? 2005 The Paleontological Society. All rights reserved.

Paleobiology

Mortality sensitivity in life-stage simulation analysis: A case study of southern sea otters

Currently, there are no generally recognized approaches for linking detailed mortality and pathology data to population-level analyses of extinction risk. We used a combination of analytical and simulation-based analyses to examine 20 years of age- and sex-specific mortality data for southern sea otters ( Enhydra lutris ), and we applied results to project the efficacy of alternative conservation strategies. Population recovery of the southern sea otter has been slow (rate of population increase λ = 1.05) compared to other recovering populations ( λ = 1.17–1.20 ), and the population declined ( λ = 0.975 ) between 1995 and 1999. Age-based Leslie matrices were developed to explore explanations for the slow recovery and recent decline in the southern sea other population. An elasticity analysis was performed to predict effects of proportional changes in stage-specific reproductive or survival rates on the rate of population increase. A life-stage simulation analysis (LSA) was developed to evaluate the impact of changing age- and cause-specific mortality rates on λ . The information used to develop these models was derived from death assemblage, pathology, and live population census data to examine the sensitivity of sea otter population growth to different sources of mortality (e.g., disease and starvation, direct human take [fisheries, gun shot, boat strike, oil pollution], mating trauma and intraspecific aggression, shark bites, and unknown). We used resampling simulations to generate random combinations of vital rates for a large number of matrix replicates and drew on these to estimate potential effects of mortality sources on population growth ( λ ). Our analyses suggest management actions that are likely and unlikely to promote recovery of the southern sea otter and more broadly indicate a methodology to better utilize cause-of-death data in conservation decision-making.

Ecological Applications

Complex trophic interactions in kelp forest ecosystems

The distributions and abundances of species and populations change almost continuously. Understanding the processes responsible is perhaps ecology’s most fundamental challenge. Kelp-forest ecosystems in southwest Alaska have undergone several phase shifts between alga- and herbivore-dominated states in recent decades. Overhunting and recovery of sea otters caused the earlier shifts. Studies focusing on these changes demonstrate the importance of top-down forcing processes, a variety of indirect food-web interactions associated with the otter-urchin-kelp trophic cascade, and the role of food-chain length in the coevolution of defense and resistance in plants and their herbivores. This system unexpectedly shifted back to an herbivore-dominated state during the 1990s, because of a sea-otter population collapse that apparently was driven by increased predation by killer whales. Reasons for this change remain uncertain but seem to be linked to the whole-sale collapse of marine mammals in the North Pacific Ocean and southern Bering Sea. We hypothesize that killer whales sequentially "fished down" pinniped and sea-otter populations after their earlier prey, the great whales, were decimated by commercial whaling. The dynamics of kelp forests in southwest Alaska thus appears to have been influenced by an ecological chain reaction that encompassed numerous species and large scales of space and time.

Bulletin of Marine Science