Search USGSSearch

Geology topics

M. T. Tinker

Publications and source records attributed to M. T. Tinker.

13 recordsLinked to original sources

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

Modeling coupled dynamics of an empirical predator-prey system to predict top predator recovery

Limited data, time, and funding lead conservation managers to make difficult choices in managing species recovery. Coupled dynamical models are relied upon for decision support, but their application to empirical predator-prey systems has generally been restricted to small, tractable species. To broaden their use in conservation decision-making, we developed a model suitable for predicting the population dynamics of a larger apex carnivore and its prey. We selected southern sea otters ( Enhydra lutris nereis ) and their primary estuarine prey as our case study and parameterized the dynamical model with data on sea otter, clam, and crab abundances; predator-prey interactions; and sea otter bioenergetics collected from Elkhorn Slough, CA. Our model, having integrated all these salient factors, was able to successfully reproduce trends in taxa abundance as well as shifts in sea otter diet composition and energy intake rates. Rich data inputs allow the model to predict population dynamics over realistic temporal scales not only for the site of data collection, but also for similar estuaries uncolonized by sea otters. Based on model projections parameterized with prey survey data from two such estuaries, Tomales Bay and Drakes Estero, we predict the sites could support over 160 sea otters and may hold potential to further species recovery. In systems with good data availability, the model has high predictive power and can provide multi-taxa projections useful for making informed management decisions.

California

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

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

Alaska

Energetic demands of immature sea otters from birth to weaning: Implications for maternal costs, reproductive behavior and population-level trends

Sea otters ( Enhydra lutris ) have the highest mass-specific metabolic rate of any marine mammal, which is superimposed on the inherently high costs of reproduction and lactation in adult females. These combined energetic demands have been implicated in the poor body condition and increased mortality of female sea otters nearing the end of lactation along the central California coast. However, the cost of lactation is unknown and currently cannot be directly measured for this marine species in the wild. Here, we quantified the energetic demands of immature sea otters across five developmental stages as a means of assessing the underlying energetic challenges associated with pup rearing that may contribute to poor maternal condition. Activity-specific metabolic rates, daily activity budgets and field metabolic rates (FMR) were determined for each developmental stage. Mean FMR of pre-molt pups was 2.29±0.81 MJ day −1 and increased to 6.16±2.46 and 7.41±3.17 MJ day −1 in post-molt pups and dependent immature animals, respectively. Consequently, daily energy demands of adult females increase 17% by 3 weeks postpartum and continue increasing to 96% above pre-pregnancy levels by the average age of weaning. Our results suggest that the energetics of pup rearing superimposed on small body size, marine living and limited on-board energetic reserves conspire to make female sea otters exceptionally vulnerable to energetic shortfalls. By controlling individual fitness, maternal behavior and pup provisioning strategies, this underlying metabolic challenge appears to be a major factor influencing current population trends in southern sea otters ( Enhydra lutris nereis ).

California

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

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

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

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

Ecology

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

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

Evolutionary Ecology Research

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

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

Deep-Sea Research Part II: Topical Studies in Ocea

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

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

Journal of Animal Ecology

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

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

Ecological Applications

Individual variation in prey selection by sea otters: Patterns, causes and implications

1. Longitudinal records of prey selection by 10 adult female sea otters on the Monterey Peninsula, California, from 1983 to 1990 demonstrate extreme inter-individual variation in diet. Variation in prey availability cannot explain these differences as the data were obtained from a common spatial-temporal area. 2. Individual dietary patterns persisted throughout our study, thus indicating that they are life-long characteristics. 3. Individual dietary patterns in sea otters appear to be transmitted along matrilines, probably by way of learning during the period of mother-young association. 4. Efficient utilization of different prey types probably requires radically different sensory/motor skills, each of which is difficult to acquire and all of which may exceed the learning and performance capacities of any single individual. This would explain the absence of generalists and inertia against switching, but not the existence of alternative specialists. 5. Such individual variation might arise in a constant environment from frequency-dependent effects, whereby the relative benefit of a given prey specialization depends on the number of other individuals utilizing that prey. Additionally, many of the sea otter's prey fluctuate substantially in abundance through time. This temporal variation, in conjunction with matrilineal transmission of foraging skills, may act to mediate the temporal dynamics of prey specializations. 6. Regardless of the exact cause, such extreme individual variation in diet has broad ramifications for population and community ecology. 7. The published literature indicates that similar patterns occur in many other species.

Journal of Animal Ecology

Killer whale predation on sea otters linking oceanic and nearshore ecosystems

After nearly a century of recovery from overhunting, sea otter populations are in abrupt decline over large areas of western Alaska. Increased killer whale predation is the likely cause of these declines. Elevated sea urchin density and the consequent deforestation of kelp beds in the nearshore community demonstrate that the otter's keystone role has been reduced or eliminated. This chain of interactions was probably initiated by anthropogenic changes in the offshore oceanic ecosystem.

Alaska