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Geology topics

M. Tim Tinker

Publications and source records attributed to M. Tim Tinker.

At least 37 records · Page 2Linked to original sources

Lactation and resource limitation affect stress responses, thyroid hormones, immune function, and antioxidant capacity of sea otters (Enhydra lutris)

Lactation is the most energetically demanding stage of reproduction in female mammals. Increased energetic allocation toward current reproduction may result in fitness costs, although the mechanisms underlying these trade‐offs are not well understood. Trade‐offs during lactation may include reduced energetic allocation to cellular maintenance, immune response, and survival and may be influenced by resource limitation. As the smallest marine mammal, sea otters (Enhydra lutris) have the highest mass‐specific metabolic rate necessitating substantial energetic requirements for survival. To provide the increased energy needed for lactation, female sea otters significantly increase foraging effort, especially during late‐lactation. Caloric insufficiency during lactation is reflected in the high numbers of maternal deaths due to End‐Lactation Syndrome in the California subpopulation. We investigated the effects of lactation and resource limitation on maternal stress responses, metabolic regulation, immune function, and antioxidant capacity in two subspecies of wild sea otters (northern: E. l. nereis and southern: E. l. kenyoni) within the California, Washington, and Alaska subpopulations. Lactation and resource limitation were associated with reduced glucocorticoid responses to acute capture stress. Corticosterone release was lower in lactating otters. Cortisol release was lower under resource limitation and suppression during lactation was only evident under resource limitation. Lactation and resource limitation were associated with alterations in thyroid hormones. Immune responses and total antioxidant capacity were not reduced by lactation or resource limitation. Southern sea otters exhibited higher concentrations of antioxidants, immunoglobulins, and thyroid hormones than northern sea otters. These data provide evidence for allocation trade‐offs during reproduction and in response to nutrient limitation but suggest self‐maintenance of immune function and antioxidant defenses despite energetic constraints. Income‐breeding strategists may be especially vulnerable to the consequences of stress and modulation of thyroid function when food resources are insufficient to support successful reproduction and may come at a cost to survival, and thereby influence population trends.

California

Defining the risk landscape in the context of pathogen pollution: Toxoplasma gondii in sea otters along the Pacific Rim

Pathogens entering the marine environment as pollutants exhibit a spatial signature driven by their transport mechanisms. The sea otter ( Enhydra lutris ), a marine animal which lives much of its life within sight of land, presents a unique opportunity to understand land–sea pathogen transmission. Using a dataset on Toxoplasma gondii prevalence across sea otter range from Alaska to California, we found that the dominant drivers of infection risk vary depending upon the spatial scale of analysis. At the population level, regions with high T. gondii prevalence had higher human population density and a greater proportion of human-dominated land uses, suggesting a strong role for population density of the felid definitive host of this parasite. This relationship persisted when a subset of data were analysed at the individual level: large-scale patterns in sea otter T. gondii infection prevalence were largely explained by individual exposure to areas of high human housing unit density, and other landscape features associated with anthropogenic land use, such as impervious surfaces and cropping land. These results contrast with the small-scale, within-region analysis, in which age, sex and prey choice accounted for most of the variation in infection risk, and terrestrial environmental features provided little variation to help in explaining observed patterns. These results underscore the importance of spatial scale in study design when quantifying both individual-level risk factors and landscape-scale variation in infection risk.

California

Gaps in kelp cover may threaten the recovery of California sea otters

Despite more than a century of federal protection, the California sea otter Enhydra lutris nereis remains threatened under the U.S. Endangered Species Act (ESA), and the population has not appreciably expanded its range in two decades. Here, we examine a novel dataset of 725 sea otter live strandings from 1984–2015 to gain insights into demographic and environmental factors underlying threats to sea otter recovery. Using multinomial logistic regression to evaluate spatiotemporal patterns of stranding causes, we demonstrate that increases in stranding rates, particularly outside the range center, are related to a substantial increase in shark bites. By contrast, trauma linked to human activities has declined dramatically, and now accounts for less than 5% of stranding cases. Within the range core, where the sea otter population seems regulated by prey availability, symptoms of energetic stress represent more than 63% of all strandings and are strongly associated with high sea otter density. Conversely, in range peripheries, the majority of strandings are caused by shark bite and neurological disease. Notably, these threats are virtually absent where nearshore habitat is characterized by at least 10% kelp canopy cover. Our analyses reveal that declining kelp cover may therefore constrain the population's spatial expansion and recovery in two key ways. Absence of kelp intensifies density‐independent threats in the range peripheries, and likely limits dispersal of reproductive females, which depend on kelp canopy for nursery habitat. These results highlight the significance of both top‐down and bottom‐up processes in population dynamics, and inform an ecosystem‐based approach to conservation planning.

California

Stability and change in kelp forest habitats at San Nicolas Island

Kelp forest communities are highly variable over space and time. Despite this complexity it has been suggested that kelp forest communities can be classified into one of 2 states: kelp dominated or sea urchin dominated. It has been further hypothesized that these represent “alternate stable states” because a site can remain in either of these states for decades before some perturbation causes a rapid shift to the other state. Our research group has maintained a subtidal community monitoring program for 38 years at San Nicolas Island consisting of twice-annual scuba-based surveys at 6 sites distributed within 4 regions around the island. Three types of perturbations are thought to be relevant to subtidal community dynamics at San Nicolas: (1) physical disturbances in the form of major storm and El Niño/Southern Oscillation (ENSO) events; (2) invertebrate diseases, which periodically decimate urchin populations; and (3) the reintroduction and subsequent increase of sea otters ( Enhydra lutris nereis ). These 3 perturbations differ in spatial and temporal specificity; physical disturbances and disease outbreaks occur periodically and could affect all 4 regions, while sea otter predation has been concentrated primarily at the West End sites over the last 15 years. The different types of perturbations and the duration of the time series at the kelp forests at San Nicolas make the data set ideal for testing the “alternate stable state” hypothesis. We use nonmetric multidimensional scaling (NMDS) to examine spatial and temporal patterns of community similarity at the 4 regions. In particular, we evaluate support for the existence of stable states, which are represented on NMDS plots as distinct spatial clusters. Community dynamics at each site approximated a biased random walk in NMDS space, with one or more basins of attraction and occasional jumps between basins. We found evidence for alternative stable states at some sites, and we show that transitions from one stable state to another may be influenced by interactions between multiple perturbations.

California

California sea otter (Enhydra lutris nereis) census results, Spring 2017

The 2017 census of southern sea otters ( Enhydra lutris nereis ) was conducted between late April and early July along the mainland coast of central California and in April at San Nicolas Island in southern California. The 3-year average of combined counts from the mainland range and San Nicolas Island was 3,186, down by 86 sea otters from the previous year. This is the second year that the official index has exceeded 3,090, the Endangered Species Act delisting threshold identified in the U.S. Fish and Wildlife Service’s Southern Sea Otter Recovery Plan (the threshold would need to be exceeded for 3 consecutive years before delisting consideration). The 5-year average trend in abundance, including both the mainland range and San Nicolas Island populations, remains positive at 2.3 percent per year. Continuing lack of growth in the range peripheries likely explains the cessation of range expansion.

California

Food abundance, prey morphology, and diet specialization influence individual sea otter tool use

Sea otters are well-known tool users, employing objects such as rocks or shells to break open invertebrate prey. We used a series of generalized linear mixed effect models to examine observational data on prey capture and tool use from 211 tagged individuals from 5 geographically defined study areas throughout the sea otter’s range in California. Our best supported model was able to explain 75% of the variation in the frequency of tool use by individual sea otters with only ecological and demographic variables. In one study area, where sea otter food resources were abundant, all individuals had similar diets focusing on preferred prey items and used tools at low to moderate frequencies (4–38% of prey captures). In the remaining areas, where sea otters were food-limited, individuals specialized on different subsets of the available prey and had a wider range of average tool-use frequency (0–98% of prey captures). The prevalence of difficult-to-access prey in individual diets was a major predictor of tool use and increased the likelihood of using tools on prey that were not difficult to access as well. Age, sex, and feeding habitat also contributed to the probability of tool use but to a smaller extent. We developed a conceptual model illustrating how food abundance, the prevalence of difficult-to-access prey, and individual diet specialization interacted to determine the likelihood that individual sea otters would use tools and considered the model’s relevance to other tool-using species.

Behavioral Ecology

Mitogenomes and relatedness do not predict frequency of tool-use by sea otters

Many ecological aspects of tool-use in sea otters are similar to those in Indo-Pacific bottlenose dolphins. Within an area, most tool-using dolphins share a single mitochondrial haplotype and are more related to each other than to the population as a whole. We asked whether sea otters in California showed similar genetic patterns by sequencing mitogenomes of 43 otters and genotyping 154 otters at 38 microsatellite loci. There were six variable sites in the mitogenome that yielded three haplotypes, one found in only a single individual. The other two haplotypes contained similar percentages (33 and 36%) of frequent tool-users and a variety of diet types. Microsatellite analyses showed that snail specialists, the diet specialist group that most frequently used tools, were no more related to each other than to the population as a whole. The lack of genetic association among tool-using sea otters compared with dolphins may result from the length of time each species has been using tools. Tool-use in dolphins appears to be a relatively recent innovation (less than 200 years) but sea otters have probably been using tools for many thousands or even millions of years.

Biology Letters

Asynchrony in craniomandibular development and growth in Enhydra lutris nereis (Carnivora: Mustelidae): Are southern sea otters born to bite?

Weaning represents a major ontogenetic dietary shift in southern sea otters ( Enhydra lutris nereis ), as juveniles must transition from depending on mother’s milk to independently processing hard-shelled invertebrates. When the skulls of juveniles have reached sufficient maturity to transition to a durophagous diet remains to be investigated. Here, we conducted a comprehensive analysis of skull development and growth and sexual dimorphism using geometric morphometric approaches in 204 southern sea otter skulls. We found that southern sea otters of both sexes exhibit dramatic changes in cranial and mandibular shape and size over ontogeny. Although the majority of these changes occur in the pup stage, full development and growth of the skull does not occur until well after weaning. We hypothesize that the slower maturation of the crania of newly weaned juveniles serves as a handicap by constraining jaw adductor muscle size, biting ability and feeding on hard-shelled prey. In our analysis of sexual dimorphism, we found significant sexual shape and size dimorphism in adult craniomandibular morphology that arose through differences in developmental and growth rates and duration. We postulate that males are selected to attain mature crania faster to presumably reach adult biting ability sooner, gaining a competitive advantage in obtaining food and in male–male agonistic interactions.

Biological Journal of the Linnean Society

Southern sea otter range expansion and habitat use in the Santa Barbara Channel, California

The re-colonization of the Santa Barbara channel by sea otters brings these ESA-listed marine mammals closer to active oil and gas production facilities, shipping lanes and naturally occurring oil and gas seeps. However, the degree to which sea otters may actually be affected by human-caused oil spills or exposure to natural oil seeps is currently unknown. Between 2012 and 2014, the U.S. Geological Survey and collaborating agencies conducted a telemetry-based study of sea otters in Santa Barbara channel, in order to provide critical information for resource managers (specifically the Bureau of Ocean Energy Management, henceforth BOEM, and the U.S. Fish and Wildlife Service, henceforth USFWS) about the spatial ecology, population status, and potential population threats to sea otters in Santa Barbara Channel, with particular reference to exposure to manmade structures and sources of oil and natural gas. Analysis of spatial monitoring data using a Bayesian-based synoptic model allowed for description of sea otter home ranges, identification of hot-spots of use, and insights into habitat selection behavior by male and female sea otters. Important findings included the deeper modal depth preferred by males versus females, strong preferences by both sexes for areas with persistent kelp canopy, and greater use of soft-sediment areas by males. The synoptic model also provided the ability to predict population-level density distribution for each sex in new habitats: by calculating the value of these probability density distributions at the known locations of natural seeps, we were able to identify those seeps with higher potential for sea otter encounters. The relative probability of occurrence at locations near to some seeps was sufficiently high (about 1% likelihood of occurrence for some of our study animals) that one would anticipate occasional encounters. Data on male and female survival, reproductive success, activity budgets, and body condition all indicated that sea otters in Santa Barbara Channel are not resource limited, and thus we would expect to see continued strong population growth in this area. However, the principal cause of death for study animals was lethal bites by white sharks, suggesting that shark bite mortality represents the single biggest threat to continued population growth in the Santa Barbara Channel.

California

Ecosystem features determine seagrass community response to sea otter foraging

Comparing sea otter recovery in California (CA) and British Columbia (BC) reveals key ecosystem properties that shape top-down effects in seagrass communities. We review potential ecosystem drivers of sea otter foraging in CA and BC seagrass beds, including the role of coastline complexity and environmental stress on sea otter effects. In BC, we find greater species richness across seagrass trophic assemblages. Furthermore, Cancer spp. crabs, an important link in the seagrass trophic cascade observed in CA, are less common. Additionally, the more recent reintroduction of sea otters, more complex coastline, and reduced environmental stress in BC seagrass habitats supports the hypotheses that sea otter foraging pressure is currently reduced there. In order to manage the ecosystem features that lead to regional differences in top predator effects in seagrass communities, we review our findings, their spatial and temporal constraints, and present a social-ecological framework for future research.

British Columbia, California

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

Predicting animal home-range structure and transitions using a multistate Ornstein-Uhlenbeck biased random walk

The home‐range concept is central in animal ecology and behavior, and numerous mechanistic models have been developed to understand home range formation and maintenance. These mechanistic models usually assume a single, contiguous home range. Here we describe and implement a simple home‐range model that can accommodate multiple home‐range centers, form complex shapes, allow discontinuities in use patterns, and infer how external and internal variables affect movement and use patterns. The model assumes individuals associate with two or more home‐range centers and move among them with some estimable probability. Movement in and around home‐range centers is governed by a two‐dimensional Ornstein‐Uhlenbeck process, while transitions between centers are modeled as a stochastic state‐switching process. We augmented this base model by introducing environmental and demographic covariates that modify transition probabilities between home‐range centers and can be estimated to provide insight into the movement process. We demonstrate the model using telemetry data from sea otters ( Enhydra lutris ) in California. The model was fit using a Bayesian Markov Chain Monte Carlo method, which estimated transition probabilities, as well as unique Ornstein‐Uhlenbeck diffusion and centralizing tendency parameters. Estimated parameters could then be used to simulate movement and space use that was virtually indistinguishable from real data. We used Deviance Information Criterion (DIC) scores to assess model fit and determined that both wind and reproductive status were predictive of transitions between home‐range centers. Females were less likely to move between home‐range centers on windy days, less likely to move between centers when tending pups, and much more likely to move between centers just after weaning a pup. These tendencies are predicted by theoretical movement rules but were not previously known and show that our model can extract meaningful behavioral insight from complex movement data.

Ecology

California sea otter ( Enhydra lutris nereis ) census results, Spring 2016

The 2016 census of southern sea otters Enhydra lutris nereis was conducted in May along the mainland coast of central California and in April at San Nicolas Island in southern California. The 3-year average of combined counts from the mainland range and San Nicolas Island was 3,272. This is the first year that the official index has exceeded 3,090, the Endangered Species Act delisting threshold suggested by the U.S. Fish and Wildlife Service (the threshold would need to be exceeded for 3 consecutive years before delisting consideration). The 5-year average trend in abundance, including both the mainland range and San Nicolas Island populations, is positive at 3.2 percent per year; however, regional trends vary, with localized declines at the southern and northern peripheries of the mainland range. The lack of population growth in the range peripheries over recent years likely explains the cessation of range expansion, with the range limits remaining almost unchanged from the previous 5 years.

California

Concentration and retention of Toxoplasma gondii surrogates from seawater by red abalone (Haliotis rufescens)

Small marine snails and abalone have been identified as high- and low-risk prey items, respectively, for exposure of threatened southern sea otters to Toxoplasma gondii , a zoonotic parasite that can cause fatal encephalitis in animals and humans. While recent work has characterized snails as paratenic hosts for T. gondii , the ability of abalone to vector the parasite has not been evaluated. To further elucidate why abalone predation may be protective against T. gondii exposure, this study aimed to determine whether: (1) abalone are physiologically capable of acquiring T. gondii ; and (2) abalone and snails differ in their ability to concentrate and retain the parasite. Abalone were exposed to T. gondii surrogate microspheres for 24 h, and fecal samples were examined for 2 weeks following exposure. Concentration of surrogates was 2–3 orders of magnitude greater in abalone feces than in the spiked seawater, and excretion of surrogates continued for 14 days post-exposure. These results indicate that, physiologically, abalone and snails can equally vector T. gondii as paratenic hosts. Reduced risk of T. gondii infection in abalone-specializing otters may therefore result from abalone's high nutritional value, which implies otters must consume fewer animals to meet their caloric needs.

Parasitology

Trade-offs between energy maximization and parental care in a central place forager, the sea otter

Between 1999 and 2014, 126 archival time–depth recorders (TDRs) were used to examine the foraging behavior of southern sea otters ( Enhydra lutris nereis ) off the coast of California, in both resource-abundant (recently occupied, low sea otter density) and resource-limited (long-occupied, high sea otter density) locations. Following predictions of foraging theory, sea otters generally behaved as energy rate maximizers. Males and females without pups employed similar foraging strategies to optimize rates of energy intake in resource-limited habitats, with some exceptions. Both groups increased overall foraging effort and made deeper, longer and more energetically costly dives as resources became limited, but males were more likely than females without pups to utilize extreme dive profiles. In contrast, females caring for young pups (≤10 weeks) prioritized parental care over energy optimization. The relative importance of parental care versus energy optimization for adult females with pups appeared to reflect developmental changes as dependent young matured. Indeed, contrary to females during the initial stages of lactation, females with large pups approaching weaning once again prioritized optimizing energy intake. The increasing prioritization of energy optimization over the course of lactation was possible due to the physiological development of pups and likely driven by the energetic deficit incurred by females early in lactation. Our results suggest that regardless of resource availability, females at the end of lactation approach a species-specific ceiling for percent time foraging and that reproductive females in the central portion of the current southern sea otter range are disproportionately affected by resource limitation.

Behavioral Ecology

Permissible Home Range Estimation (PHRE) in restricted habitats: A new algorithm and an evaluation for sea otters

Parametric and nonparametric kernel methods dominate studies of animal home ranges and space use. Most existing methods are unable to incorporate information about the underlying physical environment, leading to poor performance in excluding areas that are not used. Using radio-telemetry data from sea otters, we developed and evaluated a new algorithm for estimating home ranges (hereafter Permissible Home Range Estimation, or “PHRE”) that reflects habitat suitability. We began by transforming sighting locations into relevant landscape features (for sea otters, coastal position and distance from shore). Then, we generated a bivariate kernel probability density function in landscape space and back-transformed this to geographic space in order to define a permissible home range. Compared to two commonly used home range estimation methods, kernel densities and local convex hulls, PHRE better excluded unused areas and required a smaller sample size. Our PHRE method is applicable to species whose ranges are restricted by complex physical boundaries or environmental gradients and will improve understanding of habitat-use requirements and, ultimately, aid in conservation efforts.

PLoS ONE

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