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James A. Estes

Publications and source records attributed to James A. Estes.

At least 19 recordsLinked to original sources
USGS2025

Dynamic and context-dependent keystone species effects in kelp forests

Sea otters are an iconic keystone predator that can maintain kelp forests by preying on grazing invertebrates such as sea urchins. However, the effects of sea otters on kelp forests vary over their geographic range. Here, we analyze two 30-y datasets on kelp forest communities during the reintroduction of sea otters along the west coast of Vancouver Island, BC, Canada, and around San Nicolas Island, CA. We developed a community model to estimate species interactions as dynamic rates, varying with community state. We find evidence of a classic trophic cascade off Vancouver Island; the arrival of otters quickly led to depletion of urchins and recovery of kelp. However, this cascade was muted around San Nicolas Island, with otters, urchins, and kelp all coexisting at intermediate densities for multiple years. Our models show that this difference came from a pulse of strong otter impacts on urchins following recolonization off Vancouver Island, but not off San Nicolas Island. The mean effects of otters on urchins and urchins on kelp were not stronger in the north, indicating that interaction dynamics and not average interaction strength are key to explaining differences in community trajectories. We also find stronger multistep interaction chains in the south, arising from competitive interactions that indirectly buffered otter effects. These findings shed light on long-standing hypotheses about how interspecific interactions can alter the function of keystone species across community contexts. More broadly, we show how community change can be more accurately predicted by considering dynamic interaction strengths.

Proceedings of the National Academy of Sciences↗
USGS2021

Sea otter population collapse in southwest Alaska: Assessing ecological covariates, consequences, and causal factors

Sea otter ( Enhydra lutris ) populations in southwest Alaska declined substantially between about 1990 and the most recent set of surveys in 2015. Here we report changes in the distribution and abundance of sea otters, and covarying patterns in reproduction, mortality, body size and condition, diet and foraging behavior, food availability, health profiles, and exposure to environmental contaminants over this 25-yr period. The population decline, which resulted in densities on the order of 5% of environmental carrying capacity, ranged from Attu Island in the west to about Castle Cape (on the south side of the Alaska Peninsula) in the east. Remaining sea otters moved closer to shore and into shallow, protected habitats. Reproductive rates appeared unchanged with the decline. Although the demographic cause of the decline was clearly elevated mortality, stranded carcasses were rare or absent. The net rate of energy gain by foraging sea otters, body length and condition, and prey biomass density, all increased after the decline and varied inversely with sea otter population density beyond the area of decline. Sea otters within the area of decline showed no increases in health anomalies, disease, contaminant exposure, or abnormal gene transcription patterns as compared to animals outside the area of decline. These collective findings are inconsistent with nutritional limitation, disease, or environmental contaminants, and consistent with predation (or possibly some other density-independent factor) as the reason for the sea otter population decline. Our approach and analyses provide a broad conceptual template for thinking about and assessing the causes of wildlife population declines.

Alaska↗
USGS2020

Keystone predators govern the pathway and pace of climate impacts in a subarctic marine ecosystem

Predator loss and climate change are hallmarks of the Anthropocene yet their interactive effects are largely unknown. Here, we show that massive calcareous reefs, built slowly by the alga Clathromorphum nereostratum over centuries to millennia, are now declining because of the emerging interplay between these two processes. Such reefs, the structural base of Aleutian kelp forests, are rapidly eroding because of overgrazing by herbivores. Historical reconstructions and experiments reveal that overgrazing was initiated by the loss of sea otters, Enhydra lutris (which gave rise to herbivores capable of causing bioerosion), and then accelerated with ocean warming and acidification (which increased per capita lethal grazing by 34 to 60% compared with preindustrial times). Thus, keystone predators can mediate the ways in which climate effects emerge in nature and the pace with which they alter ecosystems.

Science↗
USGS2017

Rehabilitating sea otters: Feeling good versus being effective

This chapter examines the complexities of assessing the merits and drawbacks of wildlife rehabilitation. Wildlife rehabilitation is often costly, and the resulting benefits differ depending on whether one’s interest is in the welfare of individual animals or conserving populations. Two examples of this dilemma include the rehabilitation of oiled sea otters following the Exxon Valdez spill in Prince William Sound, Alaska, and the rehabilitation of stranded sea otter pups in central California. In the first example, substantial financial investment resulted in little or no benefits for population conservation. In the second example, the potential for population-level benefits is context dependent: in populations near carrying capacity the conservation impacts are negligible, whereas in isolated, low-density populations rehabilitation and release can be an effective conservation tool. Wildlife rehabilitation is valued by people for various reasons, but recognizing and acknowledging the difference between individual and population welfare is an important step toward effective wildlife conservation.

Book chapter↗
USGS2016

Characterizing species interactions to understand press perturbations: What is the community matrix?

The community matrix is among ecology's most important mathematical abstractions, formally encapsulating the interconnected network of effects that species have on one another's populations. Despite its importance, the term `community matrix' has been applied to matrices having differing interpretations. This has hindered the application of theory for understanding community structure and perturbation responses, particularly in the contexts of ecosystem-based management and conservation. Here we clarify the correspondence and distinctions between the Interaction matrix, the Alpha matrix and the Jacobian matrix, terms which are frequently used interchangeably and have numerous synonyms, including the term Community matrix. We illustrate how these matrices correspond to different ways of characterizing interaction strengths, how they permit insights regarding different types of press perturbations of species growth rates or abundances, and how these are related by a simple scaling relationship. Connections to additional interaction strength characterizations encapsulated by the Beta matrix, the Gamma matrix, and the Removal matrix are also discussed. Our synthesis highlights the empirical challenges that remain in using these mathematical tools to understand actual communities.

Ecology, Evolution, and Systematics↗
USGS2015

The cost of reproduction: differential resource specialization in female and male California sea otters

Intraspecific variation in behavior and diet can have important consequences for population and ecosystem dynamics. Here, we examine how differences in reproductive investment and spatial ecology influence individual diet specialization in male and female southern sea otters ( Enhydra lutris nereis ). We hypothesize that greater reproductive constraints and smaller home ranges of females lead to more pronounced intraspecific competition and increased specialization. We integrate stable carbon (δ 13 C) and nitrogen (δ 15 N) isotope analysis of sea otter vibrissae with long-term observational studies of five subpopulations in California. We define individual diet specialization as low ratios of within-individual variation (WIC) to total population niche width (TNW). We compare isotopic and observational based metrics of WIC/TNW for males and females to data on population densities, and movement patterns using both general linear and linear mixed-effects models. Consistent with our hypothesis, increasing population density is associated with increased individual diet specialization by females but not by males. Additionally, we find the amount of coastline in a sea otter’s home range positively related with individual dietary variability, with increased range span resulting in weaker specialization for both males and females. We attribute our results to sex-based differences in movement, with females needing to specialize in their small ranges to maximize energy gain, and posit that the paradigm of individual prey specialization in sea otters with increased intraspecific competition may be a pattern driven largely by females. Our work highlights a potentially broader role of sex in the mechanistic pressures promoting and maintaining diet specialization.

California↗
USGS2013

A multi-decade time series of kelp forest community structure at San Nicolas Island, California

San Nicolas Island is surrounded by broad areas of shallow subtidal habitat, characterized by dynamic kelp forest communities that undergo dramatic and abrupt shifts in community composition. Although these reefs are fished, the physical isolation of the island means that they receive less impact from human activities than most reefs in Southern California, making San Nicolas an ideal place to evaluate alternative theories about the dynamics of these communities. Here we present monitoring data from seven sampling stations surrounding the island, including data on fish, invertebrate, and algal abundance. These data are unusual among subtidal monitoring data sets in that they combine relatively frequent sampling (twice per year) with an exceptionally long time series (since 1980). Other outstanding qualities of the data set are the high taxonomic resolution captured and the monitoring of permanent quadrats and swaths where the history of the community structure at specific locations has been recorded through time. Finally, the data span a period that includes two of the strongest ENSO events on record, a major shift in the Pacific decadal oscillation, and the reintroduction of sea otters to the island in 1987 after at least 150 years of absence. These events provide opportunities to evaluate the effects of bottom-up forcing, top-down control, and physical disturbance on shallow rocky reef communities.

California↗
USGS2012

Structure and mechanism of diet specialisation: testing models of individual variation in resource use with sea otters

Studies of consumer-resource interactions suggest that individual diet specialisation is empirically widespread and theoretically important to the organisation and dynamics of populations and communities. We used weighted networks to analyze the resource use by sea otters, testing three alternative models for how individual diet specialisation may arise. As expected, individual specialisation was absent when otter density was low, but increased at high-otter density. A high-density emergence of nested resource-use networks was consistent with the model assuming individuals share preference ranks. However, a density-dependent emergence of a non-nested modular network for ‘core’ resources was more consistent with the ‘competitive refuge’ model. Individuals from different diet modules showed predictable variation in rank-order prey preferences and handling times of core resources, further supporting the competitive refuge model. Our findings support a hierarchical organisation of diet specialisation and suggest individual use of core and marginal resources may be driven by different selective pressures.

California↗
USGS2011

Gene transcription in sea otters (Enhydra lutris); development of a diagnostic tool for sea otter and ecosystem health

Gene transcription analysis for diagnosing or monitoring wildlife health requires the ability to distinguish pathophysiological change from natural variation. Herein, we describe methodology for the development of quantitative real-time polymerase chain reaction (qPCR) assays to measure differential transcript levels of multiple immune function genes in the sea otter ( Enhydra lutris ); sea otter-specific qPCR primer sequences for the genes of interest are defined. We establish a ‘reference’ range of transcripts for each gene in a group of clinically healthy captive and free-ranging sea otters. The 10 genes of interest represent multiple physiological systems that play a role in immuno-modulation, inflammation, cell protection, tumour suppression, cellular stress response, xenobiotic metabolizing enzymes, antioxidant enzymes and cell–cell adhesion. The cycle threshold (C T ) measures for most genes were normally distributed; the complement cytolysis inhibitor was the exception. The relative enumeration of multiple gene transcripts in simple peripheral blood samples expands the diagnostic capability currently available to assess the health of sea otters in situ and provides a better understanding of the state of their environment.

Molecular Ecology Resources↗
USGS2011

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↗
USGS2010

Variation in δ 13 C and δ 15 N diet–vibrissae trophic discrimination factors in a wild population of California sea otters

The ability to quantify dietary inputs using stable isotope data depends on accurate estimates of isotopic differences between a consumer (c) and its diet (d), commonly referred to as trophic discrimination factors (TDFs) and denoted by Δ c-d . At present, TDFs are available for only a few mammals and are usually derived in captive settings. The magnitude of TDFs and the degree to which they vary in wild populations is unknown. We determined δ 13 C and δ 15 N TDFs for vibrissae (i.e., whiskers), a tissue that is rapidly becoming an informative isotopic substrate for ecologists, of a wild population of sea otters for which individual diet has been quantified through extensive observational study. This is one of the very few studies that report TDFs for free-living wild animals feeding on natural diets. Trophic discrimination factors of 2.2‰ ± 0.7‰ for δ 13 C and 3.5‰ ± 0.6‰ for δ 15 N (mean ± SD) were similar to those reported for captive carnivores, and variation in individual δ 13 C TDFs was negatively but significantly related to sea urchin consumption. This pattern may relate to the lipid-rich diet consumed by most sea otters in this population and suggests that it may not be appropriate to lipid-extract prey samples when using the isotopic composition of keratinaceous tissues to examine diet in consumers that frequently consume lipid-rich foods, such as many marine mammals and seabirds. We suggest that inherent variation in TDFs should be included in isotopically based estimates of trophic level, food chain length, and mixing models used to quantify dietary inputs in wild populations; this practice will further define the capabilities and limitations of isotopic approaches in ecological studies.

Ecological Applications↗
USGS2010

Persistent organic pollutants in the blood of free-ranging sea otters (Enhydra lutris ssp.) in Alaska and California

As part of tagging and ecologic research efforts in 1997 and 1998, apparently healthy sea otters of four age-sex classes in six locations in Alaska and three in California were sampled for persistent organic pollutants (POPs) and other chemicals of ecologic or environmental concern (COECs). Published techniques for the detection of POPs (specifically Σpolychlorinated biphenyls [PCBs], ΣDDTs, Σhexachlorocyclohexanes [HCHs], Σpolycyclic aromatic hydrocarbons [PAHs], Σchlordanes [CHLs], hexachlorobenzene [HCB], dieldrin, and mirex) in the tissue of dead otters were modified for use with serum from live sea otters. Toxic equivalencies (TEQs) were calculated for POPs with proven bioactivity. Strong location effects were seen for most POPs and COECs; sea otters in California generally showed higher mean concentrations than those in Alaska. Differences in contaminant concentrations were detected among age and sex classes, with high levels frequently observed in subadults. Very high levels of ΣDDT were detected in male sea otters in Elkhorn Slough, California, where strong freshwater outflow from agricultural areas occurs seasonally. All contaminants except mirex differed among Alaskan locations; only ΣDDT, HCB, and chlorpyrifos differed within California. High levels of ΣPCB (particularly larger, more persistent congeners) were detected at two locations in Alaska where associations between elevated PCBs and military activity have been established, while higher PCB levels were found at all three locations in California where no point source of PCBs has been identified. Although POP and COEC concentrations in blood may be less likely to reflect total body burden, concentrations in blood of healthy animals may be more biologically relevant and less influenced by state of nutrition or perimortem factors than other tissues routinely sampled.

Alaska, California↗
USGS2009

Organochlorine contaminants in fishes from coastal waters west of Amukta Pass, Aleutian Islands, Alaska, USA

Organochlorines were examined in liver and stable isotopes in muscle of fishes from the western Aleutian Islands, Alaska, in relation to islands or locations affected by military occupation. Pacific cod (Gadus macrocephalus) , Pacific halibut (Hippoglossus stenolepis) , and rock greenling (Hexagrammos lagocephalus) were collected from nearshore waters at contemporary (decommissioned) and historical (World War II) military locations, as well as at reference locations. Total (Σ) polychlorinated biphenyls (PCBs) dominated the suite of organochlorine groups (ΣDDTs, Σchlordane cyclodienes, Σother cyclodienes, and Σchlorinated benzenes and cyclohexanes) detected in fishes at all locations, followed by ΣDDTs and Σchlordanes; dichlorodiphenyldi-chloroethylene ( p , p ′DDE) composed 52 to 66% of ΣDDTs by species. Organochlorine concentrations were higher or similar in cod compared to halibut and lowest in greenling; they were among the highest for fishes in Arctic or near Arctic waters. Organochlorine group concentrations varied among species and locations, but ΣPCB concentrations in all species were consistently higher at military locations than at reference locations. Moreover, all organochlorine group concentrations were higher in halibut from military locations than those from reference locations. A wide range of molecular weight organochlorines was detected at all locations, which implied regional or long-range transport and deposition, as well as local point-source contamination. Furthermore, a preponderance of higher-chlorinated PCB congeners in fishes from contemporary military islands implied recent exposure. Concentrations in all organochlorine groups increased with δ 15 N enrichment in fishes, and analyses of residual variation provided further evidence of different sources of ΣPCBs and p , p ′DDE among species and locations.

Alaska↗
USGS2009

Prey choice and habitat use drive sea otter pathogen exposure in a resource-limited coastal system

The processes promoting disease in wild animal populations are highly complex, yet identifying these processes is critically important for conservation when disease is limiting a population. By combining field studies with epidemiologic tools, we evaluated the relationship between key factors impeding southern sea otter ( Enhydra lutris nereis ) population growth: disease and resource limitation. This threatened population has struggled to recover despite protection, so we followed radio-tagged sea otters and evaluated infection with 2 disease-causing protozoal pathogens, Toxoplasma gondii and Sarcocystis neurona , to reveal risks that increased the likelihood of pathogen exposure. We identified patterns of pathogen infection that are linked to individual animal behavior, prey choice, and habitat use. We detected a high-risk spatial cluster of S. neurona infections in otters with home ranges in southern Monterey Bay and a coastal segment near San Simeon and Cambria where otters had high levels of infection with T. gondii . We found that otters feeding on abalone, which is the preferred prey in a resource-abundant marine ecosystem, had a very low risk of infection with either pathogen, whereas otters consuming small marine snails were more likely to be infected with T. gondii . Individual dietary specialization in sea otters is an adaptive mechanism for coping with limited food resources along central coastal California. High levels of infection with protozoal pathogens may be an adverse consequence of dietary specialization in this threatened species, with both depleted resources and disease working synergistically to limit recovery.

California↗
USGS2009

Otters, Marine

The otters (Mustelidae; Lutrinae) provide an exceptional perspective into the evolution of marine living by mammals. Most extant marine mammals (e.g. the cetaceans, pinnipeds, and sirenians) have been so highly modified by long periods of selection for life in the sea that they bear little resemblance to their terrestrial ancestors. Marine otters, in contrast, are more recent expatriates from freshwater habitats and some species still live in both environments. Contrasts among species within the otters, and among the otters, terrestrial mammals, and the more highly adapted pinnipeds and cetaceans provide powerful insights into mammalian adaptations to life in the sea (Estes, 1989). Among the marine mammals, sea otters ( Enhydra lutris , Fig. 1) provide the clearest understanding of consumer-induced effects on ecosystem function. This is due in part to opportunities provided by history and in part to the relative ease with which shallow coastal systems where sea otters live can be observed and studied. Although more difficult to study than sea otters, other otter species reveal the connectivity among the marine, freshwater, and terrestrial systems. These three qualities of the otters – their comparative biology, their role as predators, and their role as agents of ecosystem connectivity – are what make them interesting to marine mammalogy. The following account provides a broad overview of the comparative biology and ecology of the otters, with particular emphasis on those species or populations that live in the sea. Sea otters are features prominently, in part because they live exclusively in the sea whereas other otters have obligate associations with freshwater and terrestrial environments (Kenyon, 1969; Riedman and Estes, 1990).

Book chapter↗
USGS2008

Understanding and predicting ecological dynamics: Are major surprises inevitable

Ecological surprises, substantial and unanticipated changes in the abundance of one or more species that result from previously unsuspected processes, are a common outcome of both experiments and observations in community and population ecology. Here, we give examples of such surprises along with the results of a survey of well-established field ecologists, most of whom have encountered one or more surprises over the course of their careers. Truly surprising results are common enough to require their consideration in any reasonable effort to characterize nature and manage natural resources. We classify surprises as dynamic-, pattern-, or intervention-based, and we speculate on the common processes that cause ecological systems to so often surprise us. A long-standing and still growing concern in the ecological literature is how best to make predictions of future population and community dynamics. Although most work on this subject involves statistical aspects of data analysis and modeling, the frequency and nature of ecological surprises imply that uncertainty cannot be easily tamed through improved analytical procedures, and that prudent management of both exploited and conserved communities will require precautionary and adaptive management approaches.

Ecology↗
USGS2007

The kelp highway hypothesis: Marine ecology, the coastal migration theory, and the peopling of the Americas

In this article, a collaborative effort between archaeologists and marine ecologists, we discuss the role kelp forest ecosystems may have played in facilitating the movement of maritime peoples from Asia to the Americas near the end of the Pleistocene. Growing in cool nearshore waters along rocky coastlines, kelp forests offer some of the most productive habitats on earth, with high primary productivity, magnified secondary productivity, and three-dimensional habitat supporting a diverse array of marine organisms. Today, extensive kelp forests are found around the North Pacific from Japan to Baja California. After a break in the tropics—where nearshore mangrove forests and coral reefs are highly productive—kelp forests are also found along the Andean Coast of South America. These Pacific Rim kelp forests support or shelter a wealth of shellfish, fish, marine mammals, seabirds, and seaweeds, resources heavily used historically by coastal peoples. By about 16,000 years ago, the North Pacific Coast offered a linear migration route, essentially unobstructed and entirely at sea level, from northeast Asia into the Americas. Recent reconstructions suggest that rising sea levels early in the postglacial created a highly convoluted and island-rich coast along Beringia's southern shore, conditions highly favorable to maritime hunter-gatherers. Along with the terrestrial resources available in adjacent landscapes, kelp forests and other nearshore habitats sheltered similar suites of food resources that required minimal adaptive adjustments for migrating coastal peoples. With reduced wave energy, holdfasts for boats, and productive fishing, these linear kelp forest ecosystems may have provided a kind of “kelp highway” for early maritime peoples colonizing the New World.

Journal of Island and Coastal Archaeology↗