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M. Tim Tinker

Publications and source records attributed to M. Tim Tinker.

At least 55 records · Page 3Linked to original sources

The high cost of motherhood: End-lactation syndrome in southern sea otters ( Enhydra lutris nereis ) on the central California, USA, coast

Sea otters ( Enhydra lutris ) have exceptionally high energetic requirements, which nearly double during lactation and pup care. Thus, females are extremely vulnerable to caloric insufficiency. Despite a number of compensatory strategies, the metabolic challenge of reproduction culminates in numerous maternal deaths annually. Massive depletion of energy reserves results in a case presentation that we define as end-lactation syndrome (ELS), characterized by moderate to severe emaciation not attributable to a concurrent, independent disease process in females dying during late pup care or postweaning. We compiled detailed data for 108 adult female southern sea otters ( Enhydra lutris nereis ) examined postmortem that stranded in California, US, 2005–12, and assessed pathology, reproductive status, and the location and timing of stranding. We introduce simple, grossly apparent, standardized physical criteria to assess reproductive stage for female sea otters. We also describe ELS, examine associated risk factors, and highlight female life history strategies that likely optimize reproduction and survival. Our data suggest that females can reset both the timing and energetic demands of reproduction through fetal loss, pup abandonment, or early weaning as part of specific physiologic checkpoints during each reproductive cycle. Females appear to preload nutritionally during delayed implantation and gestation to increase fitness and reproductive success. We found that ELS was a major cause of death, affecting 56% of enrolled adult females. Peak ELS prevalence occurred in late spring, possibly reflecting the population trend toward fall/winter pupping. Increasing age and number of pregnancies were associated with a higher risk of ELS. Although the proportion of ELS females was highest in areas with dense sea otter populations, cases were recovered throughout the range, suggesting that death from ELS is associated with, but not solely caused by, population resource limitation.

California

Dramatic increase in sea otter mortality from white sharks in California

Although southern sea otters (Enhydra lutris nereis) are not considered prey for white sharks (Carcharodon carcharias), sharks do nonetheless bite sea otters. We analyzed spatial and temporal trends in shark bites on sea otters in California, assessing the frequency of shark bite wounds in 1,870 carcasses collected since 1985. The proportion of stranded sea otters having shark bites has increased sharply since 2003, and white shark bites now account for >50% of recovered carcasses. The trend was most pronounced in the southern part of the range, from Estero Bay to Point Conception, where shark bite frequency has increased eightfold. Seasonal trends were also evident: most shark-bitten carcasses are recovered in late summer and fall; however, the period of elevated shark bite frequency has lengthened. The causes of these trends are unclear, but possible contributing factors include increased white shark abundance and/or changes in white shark behavior and distribution. In particular, the spatiotemporal patterns of shark-bitten sea otters match increases in pinniped populations, and the increased availability of marine mammal prey for white sharks may have led to more sharks spending more time in nearshore waters utilized by both sea otters and pinnipeds.

California

Concentration and retention of Toxoplasma gondii oocysts by marine snails demonstrate a novel mechanism for transmission of terrestrial zoonotic pathogens in coastal ecosystems

The parasite T oxoplasma gondii is an environmentally persistent pathogen that can cause fatal disease in humans, terrestrial warm-blooded animals and aquatic mammals. Although an association between T . gondii exposure and prey specialization on marine snails was identified in threatened California sea otters, the ability of kelp-dwelling snails to transmit terrestrially derived pathogens has not been previously investigated. The objective of this study was to measure concentration and retention of T . gondii by marine snails in laboratory aquaria, and to test for natural T . gondii contamination in field-collected snails. Following exposure to T . gondii -containing seawater, oocysts were detected by microscopy in snail faeces and tissues for 10 and 3 days respectively. Nested polymerase chain reaction was also applied as a method for confirming putative T . gondii oocysts detected in snail faeces and tissues by microscopy. T oxoplasma gondii was not detected in field-collected snails. Results suggest that turban snails are competent transport hosts for T . gondii . By concentrating oocysts in faecal pellets, snails may facilitate entry of T . gondii into the nearshore marine food web. This novel mechanism also represents a general pathway by which marine transmission of terrestrially derived microorganisms can be mediated via pathogen concentration and retention by benthic invertebrates.

Environmental Microbiology

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

Timescales alter the inferred strength and temporal consistency of intraspecific diet specialization

Many populations consist of individuals that differ substantially in their diets. Quantification of the magnitude and temporal consistency of such intraspecific diet variation is needed to understand its importance, but the extent to which different approaches for doing so reflect instantaneous vs. time-aggregated measures of individual diets may bias inferences. We used direct observations of sea otter individuals ( Enhydra lutris nereis ) to assess how: (1) the timescale of sampling, (2) under-sampling, and (3) the incidence- vs. frequency-based consideration of prey species affect the inferred strength and consistency of intraspecific diet variation. Analyses of feeding observations aggregated over hourly to annual intervals revealed a substantial bias associated with time aggregation that decreases the inferred magnitude of specialization and increases the inferred consistency of individuals’ diets. Time aggregation also made estimates of specialization more sensitive to the consideration of prey frequency, which decreased estimates relative to the use of prey incidence; time aggregation did not affect the extent to which under-sampling contributed to its overestimation. Our analyses demonstrate the importance of studying intraspecific diet variation with an explicit consideration of time and thereby suggest guidelines for future empirical efforts. Failure to consider time will likely produce inconsistent predictions regarding the effects of intraspecific variation on predator–prey interactions.

Oecologia

The interaction of intraspecific competition and habitat on individual diet specialization: a near range-wide examination of sea otters

The quantification of individuality is a common research theme in the fields of population, community, and evolutionary ecology. The potential for individuality to arise is likely context-dependent, and the influence of habitat characteristics on its prevalence has received less attention than intraspecific competition. We examined individual diet specialization in 16 sea otter (Enhydra lutris) populations from southern California to the Aleutian Islands in Alaska. Because population histories, relative densities, and habitat characteristics vary widely among sites, we could examine the effects of intraspecific competition and habitat on the prevalence of individual diet specialization. Using observed diet data, we classified half of our sites as rocky substrate habitats and the other half containing a mixture of rocky and unconsolidated (soft) sediment substrates. We used stable isotope data to quantify population- and individual-level diet variation. Among rocky substrate sites, the slope [±standard error (SE)] of the positive significant relationship between the within-individual component (WIC) and total isotopic niche width (TINW) was shallow (0.23 ± 0.07) and negatively correlated with sea otter density. In contrast, the slope of the positive WIC/TINW relationship for populations inhabiting mixed substrate habitats was much higher (0.53 ± 0.14), suggesting a low degree of individuality, irrespective of intraspecific competition. Our results show that the potential for individuality to occur as a result of increasing intraspecific competition is context-dependent and that habitat characteristics, which ultimately influence prey diversity, relative abundance, and the range of skillsets required for efficient prey procurement, are important in determining when and where individual diet specialization occurs in nature.

Oecologia

Evaluating potential conservation conflicts between two listed species: Sea otters and black abalone

Population consequences of endangered species interacting as predators and prey have been considered theoretically and legally, but rarely investigated in the field. We examined relationships between spatially variable populations of a predator, the California sea otter, Enhydra lutris nereis , and a prey species, the black abalone, Haliotis cracherodii . Both species are federally listed under the Endangered Species Act and co-occur along the coast of California. We compared the local abundance and habitat distribution of black abalone at 12 sites with varying densities of sea otters. All of the populations of abalone we examined were in the geographic area currently unaffected by withering disease, which has decimated populations south of the study area. Surprisingly, our findings indicate that sea otter density is positively associated with increased black abalone density. The presence of sea otters also correlated with a shift in black abalone to habitat conferring greater refuge, which could decrease illegal human harvest. These results highlight the need for a multi-species approach to conservation management of the two species, and demonstrate the importance of using field-collected data rather than simple trophic assumptions to understand relationships between jointly vulnerable predator and prey populations.

California

The use of quantitative models in sea otter conservation

Sea otters are good indicators of ocean health. In addition, they are a keystone species, offering a stabilizing effect on ecosystem, controlling sea urchin populations that would otherwise inflict damage to kelp forest ecosystems. The kelp forest ecosystem is crucial for marine organisms and contains coastal erosion. With the concerns about the imperiled status of sea otter populations in California, Aleutian Archipelago and coastal areas of Russia and Japan, the last several years have shown growth of interest culturally and politically in the status and preservation of sea otter populations. Sea Otter Conservation brings together the vast knowledge of well-respected leaders in the field, offering insight into the more than 100 years of conservation and research that have resulted in recovery from near extinction. This publication assesses the issues influencing prospects for continued conservation and recovery of the sea otter populations and provides insight into how to handle future global changes.

Book chapter

Ecological drivers of variation in tool-use frequency across sea otter populations

Sea otters are well-known tool users, employing objects such as rocks or shells to break open hard-shelled invertebrate prey. However, little is known about how the frequency of tool use varies among sea otter populations and the factors that drive these differences. We examined 17 years of observational data on prey capture and tool use from 8 sea otter populations ranging from southern California to the Aleutian Islands in Alaska. There were significant differences in the diets of these populations as well as variation in the frequency of tool use. Sea otters at Amchitka Island, Alaska, used tools on less than 1% of dives that resulted in the capture of prey compared with approximately 16% in Monterey, California. The percentage of individuals in the population that used tools ranged from 10% to 93%. In all populations, marine snails and thick-shelled bivalves were most likely to be associated with tool use, whereas soft-bodied prey items such as worms and sea stars were the least likely. The probability that a tool would be used on a given prey type varied across populations. The morphology of the prey item being handled and the prevalence of various types of prey in local diets were major ecological drivers of tool use: together they accounted for about 64% of the variation in tool-use frequency among populations. The remaining variation may be related to changes in the relative costs and benefits to an individual otter of learning to use tools effectively under differing ecological circumstances.

Alaska, California

Testing the nutritional-limitation, predator-avoidance, and storm-avoidance hypotheses for restricted sea otter habitat use in the Aleutian Islands, Alaska

Sea otters ( Enhydra lutris) inhabiting the Aleutian Islands have stabilized at low abundance levels following a decline and currently exhibit restricted habitat-utilization patterns. Possible explanations for restricted habitat use by sea otters can be classified into two fundamentally different processes, bottom-up and top-down forcing. Bottom-up hypotheses argue that changes in the availability or nutritional quality of prey resources have led to the selective use of habitats that support the highest quality prey. In contrast, top-down hypotheses argue that increases in predation pressure from killer whales have led to the selective use of habitats that provide the most effective refuge from killer whale predation. A third hypothesis suggests that current restricted habitat use is based on a need for protection from storms. We tested all three hypotheses for restricted habitat use by comparing currently used and historically used sea otter foraging locations for: (1) prey availability and quality, (2) structural habitat complexity, and (3) exposure to prevailing storms. Our findings suggest that current use is based on physical habitat complexity and not on prey availability, prey quality, or protection from storms, providing further evidence for killer whale predation as a cause for restricted sea otter habitat use in the Aleutian Islands.

Alaska

An online database for informing ecological network models: http://kelpforest.ucsc.edu

Ecological network models and analyses are recognized as valuable tools for understanding the dynamics and resiliency of ecosystems, and for informing ecosystem-based approaches to management. However, few databases exist that can provide the life history, demographic and species interaction information necessary to parameterize ecological network models. Faced with the difficulty of synthesizing the information required to construct models for kelp forest ecosystems along the West Coast of North America, we developed an online database ( http://kelpforest.ucsc.edu/ ) to facilitate the collation and dissemination of such information. Many of the database's attributes are novel yet the structure is applicable and adaptable to other ecosystem modeling efforts. Information for each taxonomic unit includes stage-specific life history, demography, and body-size allometries. Species interactions include trophic, competitive, facilitative, and parasitic forms. Each data entry is temporally and spatially explicit. The online data entry interface allows researchers anywhere to contribute and access information. Quality control is facilitated by attributing each entry to unique contributor identities and source citations. The database has proven useful as an archive of species and ecosystem-specific information in the development of several ecological network models, for informing management actions, and for education purposes (e.g., undergraduate and graduate training). To facilitate adaptation of the database by other researches for other ecosystems, the code and technical details on how to customize this database and apply it to other ecosystems are freely available and located at the following link ( https://github.com/kelpforest-cameo/databaseui ).

PLoS ONE

Effects of wildfire on sea otter (Enhydra lutris) gene transcript profiles

Wildfires have been shown to impact terrestrial species over a range of temporal scales. Little is known, however, about the more subtle toxicological effects of wildfires, particularly in downstream marine or downwind locations from the wildfire perimeter. These down-current effects may be just as substantial as those effects within the perimeter. We used gene transcription technology, a sensitive indicator of immunological perturbation, to study the effects of the 2008 Basin Complex Fire on the California coast on a sentinel marine species, the sea otter ( Enhydra lutris ). We captured sea otters in 2008 (3 mo after the Basin Complex Fire was controlled) and 2009 (15 mo after the Basin Complex Fire was controlled) in the adjacent nearshore environment near Big Sur, California. Gene responses were distinctly different between Big Sur temporal groups, signifying detoxification of PAHs, possible associated response to potential malignant transformation, and suppression of immune function as the primary responses of sea otters to fire in 2008 compared to those captured in 2009. In general, gene transcription patterns in the 2008 sea otters were indicative of molecular reactions to organic exposure, malignant transformation, and decreased ability to respond to pathogens that seemed to consistent with short-term hydrocarbon exposure.

California

Sea otters are recolonizing southern California in fits and starts

After near extinction as a result of the fur trade in the 1700s and 1800s, the southern sea otter slowly reoccupied the core of its range in central California. Range expansion beyond central California is seen as key to full recovery of otters, but the rate of expansion has been sporadic, raising concerns about habitat quality in southern California. To describe the range expansion of sea otters from central into southern California, we used skiff surveys, aerial surveys, and archival time-depth recorders from 2004 to 2013. These observations show that range expansion began when male otters swam southeast of Point Conception (Cojo Anchorage), perhaps to seek refuge from bad weather and to feed on unexploited resources. After several years of seasonal use by male groups, females began to use the area, leading to reproduction and a secondary increase in abundance. In contrast, a second male group that moved farther down the coast to Coal Oil Point stalled and retreated. Such range expansion and contraction can be explained by the social nature of sea otters, which acts to slow dispersal away from groups. Otter densities at Cojo Anchorage are now approaching equilibrium levels reported for central California. As in central California, otters rested in and near kelp forest habitat, but used deeper water for foraging. Together, these observations suggest habitat in the Santa Barbara Channel can still support sea otters, but range expansion of otters into southern California will be episodic due to social dynamics.

California

Recovery of a top predator mediates negative eutrophic effects on seagrass

A fundamental goal of the study of ecology is to determine the drivers of habitat-forming vegetation, with much emphasis given to the relative importance to vegetation of “bottom-up” forces such as the role of nutrients and “top-down” forces such as the influence of herbivores and their predators. For coastal vegetation (e.g., kelp, seagrass, marsh, and mangroves) it has been well demonstrated that alterations to bottom-up forcing can cause major disturbances leading to loss of dominant vegetation. One such process is anthropogenic nutrient loading, which can lead to major changes in the abundance and species composition of primary producers, ultimately affecting important ecosystem services. In contrast, much less is known about the relative importance of apex predators on coastal vegetated ecosystems because most top predator populations have been depleted or lost completely. Here we provide evidence that an unusual four-level trophic cascade applies in one such system, whereby a top predator mitigates the bottom-up influences of nutrient loading. In a study of seagrass beds in an estuarine ecosystem exposed to extreme nutrient loading, we use a combination of a 50-y time series analysis, spatial comparisons, and mesocosm and field experiments to demonstrate that sea otters ( Enhydra lutris ) promote the growth and expansion of eelgrass ( Zostera marina ) through a trophic cascade, counteracting the negative effects of agriculturally induced nutrient loading. Our results add to a small but growing body of literature illustrating that significant interactions between bottom-up and top-down forces occur, in this case with consequences for the conservation of valued ecosystem services provided by seagrass.

California

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

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

Tools for quantifying isotopic niche space and dietary variation at the individual and population level.

Ecologists are increasingly using stable isotope analysis to inform questions about variation in resource and habitat use from the individual to community level. In this study we investigate data sets from 2 California sea otter ( Enhydra lutris nereis ) populations to illustrate the advantages and potential pitfalls of applying various statistical and quantitative approaches to isotopic data. We have subdivided these tools, or metrics, into 3 categories: IsoSpace metrics, stable isotope mixing models, and DietSpace metrics. IsoSpace metrics are used to quantify the spatial attributes of isotopic data that are typically presented in bivariate (e.g., δ 13 C versus δ 15 N) 2-dimensional space. We review IsoSpace metrics currently in use and present a technique by which uncertainty can be included to calculate the convex hull area of consumers or prey, or both. We then apply a Bayesian-based mixing model to quantify the proportion of potential dietary sources to the diet of each sea otter population and compare this to observational foraging data. Finally, we assess individual dietary specialization by comparing a previously published technique, variance components analysis, to 2 novel DietSpace metrics that are based on mixing model output. As the use of stable isotope analysis in ecology continues to grow, the field will need a set of quantitative tools for assessing isotopic variance at the individual to community level. Along with recent advances in Bayesian-based mixing models, we hope that the IsoSpace and DietSpace metrics described here will provide another set of interpretive tools for ecologists.

Journal of Mammalogy

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