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James A. Estes

Publications and source records attributed to James A. Estes.

At least 37 records · Page 2Linked to original sources

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↗

Sea otter mortality from the Exxon Valdez spill: Evaluation of an estimate from boat-based surveys

The Exxon Valadez oil spill killed large numbers of sea otters ( Enhydra lutris ) in Prince William Sound (PWS), Alaska, where the spill originated, as well as within the oil’s path along the coasts of the Kenai Peninsula and Kodiak Island. Sea otters attracted particular attention after the spill due to their public appeal (Batten 1990) and known vulnerability to oil (Costa and Kooyman 1982, Siniff et al. 1982, Davis et al. 1988, Williams et al. 1988). Extensive efforts were made to rescue and rehabilitate oiled otters at a resulting cost of ca. $80,000 per animal (Estes 1991). The argument that otters lost in the spill should be similarly valued made the estimate of mortality a matter of substantial concern to those involved in spill-related litigation or out-of-court settlements.

Alaska↗

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. ?? 2006 by the Ecological Society of America.

Ecological Applications↗

Catastrophe, recovery and range limitation in NE Pacific kelp forests: a large-scale perspective

The 1997–98 El Niño was one of the strongest on record and resulted in widespread losses of the giant kelp Macrocystis pyrifera (Agardh) along the west coast of North America. Drawing on a rich history of studies that have shown abnormally large waves and warm nutrient-poor water associated with El Niños to negatively impact giant kelp populations at some locations in southern and Baja California, we examined (1) how these impacts scale up when considered across the species’ geographic range in the NE Pacific Ocean and (2) if these impacts are generalizable over broad spatial scales. Working at 56 sites in 14 study locations over a 3 yr period (1997 to 2000), we examined how giant kelp populations were impacted by and recovered following the 1997–98 El Niño over a ~1500 km span along the west coast of North America. Our results indicate that while nearly all giant kelp disappeared from the southern one-third of the species’ range along the coast of Baja California, Mexico, and heavy losses occurred throughout the central one-third of the species’ range in southern California, USA, only minor impacts were observed throughout the northern one-third of the species’ range in central California. Further, although highly variable among regions, these impacts were similar and generalizable among locations within each region. Our results also suggest that, as has been observed in local-scale studies, this large-scale variability in giant kelp mortality was driven by large-scale patterns in ocean temperature (nutrient concentration) and wave intensity. Recovery following El Niño, in contrast, was variable at multiple spatial scales and although not directly tested here, presumably influenced by numerous factors such as proximity to upwelling areas, competition with other algae, grazing, and propagule availability. Further, variability in the rates of recovery among locations resulted in a generally slow recovery of giant kelp throughout most of Baja California, and residual large-scale impacts of the El Niño were still evident 2 yr after the El Niño ended. As global climate change may lead to increases in the frequency and intensity of El Niños, our findings have broad implications for the ways in which ecosystems might be expected to respond to them and provide a measure by which their impacts to giant kelp ecosystems may be compared among events.

Pacific Ocean↗

Foraging ecology

Longitudinal foraging data collected from 60 sea otters implanted with VHF radio transmitters at two study sites in Central California over a three-year period demonstrated even greater individual dietary specialization than in previous studies, with only 54% dietary overlap between individuals and the population. Multivariate statistical analyses indicated that individual diets could be grouped into three general "diet types" representing distinct foraging specializations. Type 1 specialists consumed large size prey but had low dive efficiency, Type 2 specialists consumed small to medium size prey with high dive efficiency, and Type 3 specialists consumed very small prey (mainly snails) with very high dive efficiency. The mean rate of energy gain for the population as a whole was low when compared to other sea otter populations in Alaska but showed a high degree of within- and betweenindividual variation, much of which was accounted for by the three foraging strategies. Type 1 specialists had the highest mean energy gain but also the highest withinindividual variance in energy gain. Type 2 specialists had the lowest mean energy gain but also the lowest variance. Type 3 specialists had an intermediate mean and variance. All three strategies resulted in very similar probabilities of exceeding a critical rate of energy gain on any given day. Correlational selection may help maintain multiple foraging strategies in the population: a fitness surface (using mean rate of energy gain as a proxy for fitness) fit to the first two principal components of foraging behavior suggested that the three foraging strategies occupy separate fitness peaks. Food limitation is likely an important ultimate factor restricting population growth in the center of the population’s range in California, although the existence of alternative foraging strategies results in different impacts of food limitation on individuals and thus may obscure expected patterns of density dependence.

Book chapter↗

Spatial and temporal variation in sea otter demography

1) Better information on historical and current population dynamics is central to understanding patterns of growth and decline in the California sea otter population. We developed a maximum likelihood-based analytical method to estimate historical age/sex specific vital rates as well as spatial and temporal variation in vital rates from longitudinal databases on population census numbers and the age-structure of salvaged carcasses. 2) We estimated current demographic parameters by conducting a mark-recapture study, measuring survival and reproduction of 115 radio-tagged individuals between 2001 and 2004. These current estimates were compared to estimates from a similar study of radiotagged otters conducted in the mid-eighties. 3) Together, these two approaches indicated that survival has decreased substantially between the early 1990s and the present and is lowest in the north-central portion of the population's range. 4) The greatest decrease in survival was for adult females (≥ 4 years of age). Variation in the survival of this age/sex class is primarily responsible for regulating population growth and driving population trends.

Book chapter↗

Re-wilding North America

A plan to restore animals that disappeared 13,000 years ago from Pleistocene North America offers an alternative conservation strategy for the twenty-first century, argue Josh Donlan and colleagues.

Nature↗

Response from Soulé and Estes

The letter from Kimberly Heiman points out the applicability of our framework to nonnative invasive species. Indeed, many alien species are changing the sign and increasing the amplitude of strong interactions in contemporary ecosystems, in some cases catastrophically. Invasive exotic species require much more attention at the policy and administrative levels.

BioScience↗

Determinants of reproductive costs in the long-lived Black-legged Kittiwake: A multiyear experiment

We studied reproductive costs of Black-legged Kittiwakes ( Rissa tridactyla ) in Prince William Sound, Alaska (USA) by removing entire clutches from randomly selected nests over four successive years, and then contrasting survival and fecundity of adults from manipulated and unmanipulated nests in each subsequent year. To elucidate mechanisms that lead to the expression of reproductive costs, we simultaneously characterized several behavioral and physiological parameters among adults in the two treatment groups. We also examined naturally nonbreeding adults that previously bred to determine their survival and future nonbreeding probabilities. Food availability varied during the study, being generally poor at the onset, and improving in later years. Adult nest attendance and body condition (assessed late in the chick- rearing period) varied accordingly among years, and between adults raising chicks and adults that had their eggs removed. Adults from unmanipulated nests incurred significant survival costs in all years, although fecundity costs were strongly expressed in only one of four years. Neither survival nor fecundity costs were strongly influenced by body condition or food availability, and no difference in reproductive costs was detected between the sexes. Although unmanipulated breeders survived at lower rates than manipulated breeders due to costs of reproduction, their survival rates were elevated compared to those of natural nonbreeders, presumably due to differences in individual ability. These findings indicate that models of adult survival must consider not only an organism's reproductive state, but also the factors that lead to that state. Although body condition appeared to be weakly related to survival, it was insufficient to explain the full magnitude of survival costs observed. We suggest that other parameters that were found to differ between treatment groups (e.g., rates of energy turnover, baseline levels of stress, and patterns of allocating body reserves) may be important mechanistic determinants of reproductive costs in kittiwakes, and potentially other long-lived species. Future efforts should move beyond simple assessments of body condition and toward more integrated measures of physiological condition when attempting to identify factors that influence how long-lived species balance the costs and benefits of reproduction.

Alaska↗

Status of sea otter populations in southcentral and southeast Alaska, 2002-2003

During the years 2002-2004 estimated sea otter population sizes were calculated for Southeast Alaska, Prince William Sound, and the Kenai Peninsula and Cook Inlet regions of Alaska. Aerial surveys were conducted by a single observer from a float-equipped Bellanca Scout fixed-wing aircraft flying at 91 m altitude and 65 mph. The surveys followed protocols written by Bodkin and Udevitz (1999). The survey design consisted of systematic sampling of 400 m wide transects that were uniformly placed throughout the survey area. Selection and sampling of transects was proportional to expected sea otter abundance, with most effort taking place in transects over waters 0-40 m in depth. Intensive searches were periodically conducted within transects to estimate the proportion of sea otters not detected on strips. To obtain an adjusted population size estimate, strip counts are adjusted for the area not surveyed and by a correction factor.

Alaska↗

Life history plasticity and population regulation in sea otters

We contrasted body condition, and age-specific reproduction and mortality between a growing population of sea otters ( Enhydra lutris ) at Kodiak Island and a high-density near-equilibrium population at Amchitka Island, Alaska. We obtained data from marked individuals, population surveys, and collections of beach-cast carcasses. Mass:length ratios indicated that females (but not males) captured in 1992 at Amchitka were in poorer condition than those captured at Kodiak in 1986–1987. In 1993, the condition of females at Amchitka improved in apparent response to two factors: (1) an episodic influx of Pacific smooth lumpsuckers, Aptocyclus ventricocus , from the epi-pelagic zone, which otters consumed; and (2) an increase in the otters’ benthic invertebrate prey resulting from declining otter numbers. Reproductive rates varied with age (0.37 [CI=0.21 to 0.53] births female −1 yr −1 for 2–3-yr-olds, and 0.83 [CI=0.69 to 0.90] for females ≥4 yr old), and were similar at both areas. Weaning success (pups surviving to ≥120 d), in contrast, was almost 50% lower at Amchitka than at Kodiak and for females ≥4 yr of age was 0.52 (CI=0.38 to 0.66) vs 0.94 (CI=0.75 to 0.99), respectively. Sixty-two percent of the preweaning pup losses at Amchitka occurred within a month of parturition and 79% within two months. Postweaning survival was also low at Amchitka as only 18% of instrumented pups were known to be alive one year after mother-pup separation. Adult survival rates appeared similar at Amchitka and Kodiak. Factors affecting survival early in life thus are a primary demographic mechanism of population regulation in sea otters. By maintaining uniformly high reproductive rates over time and limiting investment in any particular reproductive event, sea otters can take advantage of unpredictable environmental changes favorable to pup survival. This strategy is consistent with predictions of “bet-hedging” life history models.

Oikos↗

Kelp forest ecosystems: Biodiversity, stability, resilience and future

Kelp forests are phyletically diverse, structurally complex and highly productive components of coldwater rocky marine coastlines. This paper reviews the conditions in which kelp forests develop globally and where, why and at what rate they become deforested. The ecology and long archaeological history of kelp forests are examined through case studies from southern California, the Aleutian Islands and the western North Atlantic, well-studied locations that represent the widest possible range in kelp forest biodiversity. Global distribution of kelp forests is physiologically constrained by light at high latitudes and by nutrients, warm temperatures and other macrophytes at low latitudes. Within mid-latitude belts (roughly 40–60° latitude in both hemispheres) well-developed kelp forests are most threatened by herbivory, usually from sea urchins. Overfishing and extirpation of highly valued vertebrate apex predators often triggered herbivore population increases, leading to widespread kelp deforestation. Such deforestations have the most profound and lasting impacts on species-depauperate systems, such as those in Alaska and the western North Atlantic. Globally urchin-induced deforestation has been increasing over the past 2–3 decades. Continued fishing down of coastal food webs has resulted in shifting harvesting targets from apex predators to their invertebrate prey, including kelp-grazing herbivores. The recent global expansion of sea urchin harvesting has led to the widespread extirpation of this herbivore, and kelp forests have returned in some locations but, for the first time, these forests are devoid of vertebrate apex predators. In the western North Atlantic, large predatory crabs have recently filled this void and they have become the new apex predator in this system. Similar shifts from fish- to crab-dominance may have occurred in coastal zones of the United Kingdom and Japan, where large predatory finfish were extirpated long ago. Three North American case studies of kelp forests were examined to determine their long history with humans and project the status of future kelp forests to the year 2025. Fishing impacts on kelp forest systems have been both profound and much longer in duration than previously thought. Archaeological data suggest that coastal peoples exploited kelp forest organisms for thousands of years, occasionally resulting in localized losses of apex predators, outbreaks of sea urchin populations and probably small-scale deforestation. Over the past two centuries, commercial exploitation for export led to the extirpation of sea urchin predators, such as the sea otter in the North Pacific and predatory fishes like the cod in the North Atlantic. The large-scale removal of predators for export markets increased sea urchin abundances and promoted the decline of kelp forests over vast areas. Despite southern California having one of the longest known associations with coastal kelp forests, widespread deforestation is rare. It is possible that functional redundancies among predators and herbivores make this most diverse system most stable. Such biodiverse kelp forests may also resist invasion from non-native species. In the species-depauperate western North Atlantic, introduced algal competitors carpet the benthos and threaten future kelp dominance. There, other non-native herbivores and predators have become established and dominant components of this system. Climate changes have had measurable impacts on kelp forest ecosystems and efforts to control the emission of greenhouse gasses should be a global priority. However, overfishing appears to be the greatest manageable threat to kelp forest ecosystems over the 2025 time horizon. Management should focus on minimizing fishing impacts and restoring populations of functionally important species in these systems.

Environmental Conservation↗

Sea otter population declines in the Aleutian Archipelago

Sea otter ( Enhydra lutris ) populations were exploited to near extinction and began to recover after the cessation of commercial hunting in 1911. Remnant colonies of sea otters in the Aleutian archipelago were among the first to recover; they continued to increase through the 1980s but declined abruptly during the 1990s. We conducted an aerial survey of the Aleutian archipelago in 2000 and compared results with similar surveys conducted in 1965 and 1992. The number of sea otters counted decreased by 75% between 1965 and 2000; 88% for islands at equilibrial density in 1965. The population decline likely began in the mid-1980s and declined at a rate of 17.5%/year in the 1990s. The minimal population estimate was 8,742 sea otters in 2000. The population declined to a uniformly low density in the archipelago, suggesting a common and geographically widespread cause. These data are in general agreement with the hypothesis of increased predation on sea otters. These data chronicle one of the most widespread and precipitous population declines for a mammalian carnivore in recorded history.

Alaska↗

Clinical pathology and assessment of pathogen exposure in southern and Alaskan sea otters

The southern sea otter ( Enhydra lutris nereis ) population in California (USA) and the Alaskan sea otter ( E. lutris kenyoni ) population in the Aleutian Islands (USA) chain have recently declined. In order to evaluate disease as a contributing factor to the declines, health assessments of these two sea otter populations were conducted by evaluating hematologic and/or serum biochemical values and exposure to six marine and terrestrial pathogens using blood collected during ongoing studies from 1995 through 2000. Samples from 72 free-ranging Alaskan, 78 free-ranging southern, and (for pathogen exposure only) 41 debilitated southern sea otters in rehabilitation facilities were evaluated and compared to investigate regional differences. Serum chemistry and hematology values did not indicate a specific disease process as a cause for the declines. Statistically significant differences were found between free-ranging adult southern and Alaskan population mean serum levels of creatinine kinase, alkaline phosphatase, alanine aminotransferase, aspartate aminotransferase, calcium, cholesterol, creatinine, glucose, phosphorous, total bilirubin, blood urea nitrogen, and sodium. These were likely due to varying parasite loads, contaminant exposures, and physiologic or nutrition statuses. No free-ranging sea otters had signs of disease at capture, and prevalences of exposure to calicivirus, Brucella spp., and Leptospira spp. were low. The high prevalence (35%) of antibodies to Toxoplasma gondii in free-ranging southern sea otters, lack of antibodies to this parasite in Alaskan sea otters, and the pathogen's propensity to cause mortality in southern sea otters suggests that this parasite may be important to sea otter population dynamics in California but not in Alaska. The evidence for exposure to pathogens of public health importance (e.g., Leptospira spp., T. gondii ) in the southern sea otter population, and the naïveté of both populations to other pathogens (e.g., morbillivirus and Coccidiodes immitis ) may have important implications for their management and recovery.

Journal of Wildlife Diseases↗

Activity patterns and time budgets of the declining sea otter population at Amchitka Island, Alaska

Time budgets of predators may reflect population status if time spent foraging varies with local prey abun- dance. We assumed that the sea otter (Enhydra lutris) population at Amchitka Island, Alaska, USA, had been at equilibrium since the early 1960s and collected time budgets of otters to be used to represent future conditions of currently expanding sea otter populations. We used radiotelemetry to monitor activity-time budgets of otters from August 1992 to March 1994. Sea otter activity was directly linked to sex, age, weather condition, season, and time of day. Sea otters differed in percent time foraging among cohorts but not within cohorts. Percent time foraging ranged from 21% for females with very young (≤ 3weeks of age) dependent pups to 52% for females with old (≥10 weeks of age) pups. Otters foraged more and hauled out more as local sea conditions worsened. Adult males spent less time foraging during winter and spring, consistent with seasonal changes in prey selection. Time spent for- aging was similar to that reported for otters in California and an established population in Prince William Sound, Alaska, but greater than that of otters in recently established populations in Oregon and Alaska. Despite current evidence indicating that the population was in decline during our study, we were unable to recognize this change using time budgets. Our results illustrate the importance of stratifying analyses of activity patterns by age and sex cohorts and the complexity inherent in comparisons of behavioral data between different populations relying on distinct prey bases.

Journal of Wildlife Management↗

Otters

The otters (Mustelidae; Lutrinae) provide a unique look into the evolution of marine living by mammals. This is because most extant marine mammals have been so highly modified by long periods of selection for life in the sea that they bare little resemblance to their terrestrial ancestors. Marine otters, by contrast, are recent expatriates from terrestrial and freshwater habitats, and some species still live in both environments. Contrasts within this group, and among the otters, terrestrial mammals, and the more highly adapted pinnipeds and cetaceans potentially offer deep insight into mammalian adaptations to life in the sea. Among the marine mammals, sea otters also provide the clearest understanding of predation and ocean ecosystem function. This is due in part to serendipitous opportunities provided by history and in part by the relative ease with which shallow coastal systems where sea otters live can be observed and studied. These two qualities of the otters are what make them interesting to marine mammalogy. Thus, our contribution to this volume on the marine mammals is built around these themes.

Book chapter↗