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At least 163 records · Page 9Linked to original sources

Improved arrival-date estimates of Arctic-breeding Dunlin ( Calidris alpina arcticola )

The use of stable isotopes in animal ecology depends on accurate descriptions of isotope dynamics within individuals. The prevailing assumption that laboratory-derived isotopic parameters apply to free-living animals is largely untested. We used stable carbon isotopes (δ 13 C) in whole blood from migratory Dunlin ( Calidris alpina arcticola ) to estimate an in situ turnover rate and individual diet-switch dates. Our in situ results indicated that turnover rates were higher in free-living birds, in comparison to the results of an experimental study on captive Dunlin and estimates derived from a theoretical allometric model. Diet-switch dates from all 3 methods were then used to estimate arrival dates to the Arctic; arrival dates calculated with the in situ turnover rate were later than those with the other turnover-rate estimates, substantially so in some cases. These later arrival dates matched dates when local snow conditions would have allowed Dunlin to settle, and agreed with anticipated arrival dates of Dunlin tracked with light-level geolocators. Our study presents a novel method for accurately estimating arrival dates for individuals of migratory species in which return dates are difficult to document. This may be particularly appropriate for species in which extrinsic tracking devices cannot easily be employed because of cost, body size, or behavioral constraints, and in habitats that do not allow individuals to be detected easily upon first arrival. Thus, this isotopic method offers an exciting alternative approach to better understand how species may be altering their arrival dates in response to changing climatic conditions.

The Auk

Effects of microhabitat and large-scale land use on stream salamander occupancy in the coalfields of Central Appalachia

Large-scale coal mining practices, particularly surface coal extraction and associated valley fills as well as residential wastewater discharge, are of ecological concern for aquatic systems in central Appalachia. Identifying and quantifying alterations to ecosystems along a gradient of spatial scales is a necessary first-step to aid in mitigation of negative consequences to aquatic biota. In central Appalachian headwater streams, apart from fish, salamanders are the most abundant vertebrate predator that provide a significant intermediate trophic role linking aquatic and terrestrial food webs. Stream salamander species are considered to be sensitive to aquatic stressors and environmental alterations, as past research has shown linkages among microhabitat parameters, large-scale land use such as urbanization and logging, and salamander abundances. However, there is little information examining these relationships between environmental conditions and salamander occupancy in the coalfields of central Appalachia. In the summer of 2013, 70 sites (sampled two to three times each) in the southwest Virginia coalfields were visited to collect salamanders and quantify stream and riparian microhabitat parameters. Using an information-theoretic framework, effects of microhabitat and large-scale land use on stream salamander occupancy were compared. The findings indicate that Desmognathus spp. occupancy rates are more correlated to microhabitat parameters such as canopy cover than to large-scale land uses. However, Eurycea spp . occupancy rates had a strong association with large-scale land uses, particularly recent mining and forest cover within the watershed. These findings suggest that protection of riparian habitats is an important consideration for maintaining aquatic systems in central Appalachia. If this is not possible, restoration riparian areas should follow guidelines using quick-growing tree species that are native to Appalachian riparian areas. These types of trees would rapidly establish a canopy cover, stabilize the soil, and impede invasive plant species which would, in turn, provide high-quality refuges for stream salamanders.

Journal of Ecology and the Natural Environment

Defining the next generation modeling of coastal ecotone dynamics in response to global change

Coastal ecosystems are especially vulnerable to global change; e.g., sea level rise (SLR) and extreme events. Over the past century, global change has resulted in salt-tolerant (halophytic) plant species migrating into upland salt-intolerant (glycophytic) dominated habitats along major rivers and large wetland expanses along the coast. While habitat transitions can be abrupt, modeling the specific drivers of abrupt change between halophytic and glycophytic vegetation is not a simple task. Correlative studies, which dominate the literature, are unlikely to establish ultimate causation for habitat shifts, and do not generate strong predictive capacity for coastal land managers and climate change adaptation exercises. In this paper, we first review possible drivers of ecotone shifts for coastal wetlands, our understanding of which has expanded rapidly in recent years. Any exogenous factor that increases growth or establishment of halophytic species will favor the ecotone boundary moving upslope. However, internal feedbacks between vegetation and the environment, through which vegetation modifies the local microhabitat (e.g., by changing salinity or surface elevation), can either help the system become resilient to future changes or strengthen ecotone migration. Following this idea, we review a succession of models that have provided progressively better insight into the relative importance of internal positive feedbacks versus external environmental factors. We end with developing a theoretical model to show that both abrupt environmental gradients and internal positive feedbacks can generate the sharp ecotonal boundaries that we commonly see, and we demonstrate that the responses to gradual global change (e.g., SLR) can be quite diverse.

Ecological Modelling

The confluences of ideas leading to, and the flow of ideas emerging from, individual-based modeling of riverine fishes

In this review article, we trace the history of events leading to the development of individual-based models (IBMs) to represent aquatic organisms in rivers and streams. As a metaphor, we present this history as a series of confluences between individual scientists (tributaries) sharing ideas. We describe contributions of these models to science and management. One iconic feature of river IBMs is the linkage between flow and the physical habitat experienced by individual animals, and the first model that focused on this linkage is briefly described. We continue by reviewing the contributions of riverine IBMs to eight broad areas of scientific inquiry. The first four areas include research to understand 1) the effects of flow regimes on fish populations, 2) species interactions (e.g., size-mediated competition and predation), 3) fish movement and habitat selection, and 4) contaminant and water quality impacts on populations. Next, we review research using IBMs 5) to guide conservation biology of imperiled taxa through population viability analysis, including research 6) to understand river fragmentation by dams and reconnection, 7) to understand genetic outcomes for riverine metapopulations, and 8) to anticipate the future effects of temperature and climate change. This rich body of literature has contributed to both theoretical insights (e.g., about animal behavior and life history) and applied insights (e.g., population-level effects of flow regimes, temperature, and the effects of hydropower and other industries that share rivers with aquatic biota). We finish by exploring promising branches that lie ahead in the braided, downstream channel that represents future river modeling research.

Ecological Modelling

The influence of food abundance, food dispersion and habitat structure on territory selection and size of an Afrotropical terrestrial insectivore

Most tropical insectivorous birds, unlike their temperate counterparts, hold and defend a feeding and breeding territory year-around. However, our understanding of ecological factors influencing territory selection and size in tropical insectivores is limited. Here we examine three prominent hypotheses relating food abundance, food dispersion (spatial arrangement of food items), and habitat structure to territoriality in the Usambara Thrush Turdus roehli . We first compared leaf-litter macro-invertebrate abundance and dispersion, and habitat structure between territories and random sites. We then examined the relation between these same ecological factors and territory size. Invertebrate abundance and dispersion were sparsely and evenly distributed across our study system and did not vary between territories and random sites. In contrast, habitat structure did vary between territories and random sites indicating the Usambara Thrush selects territories with open understorey and closed overstorey habitat. Invertebrate abundance and dispersion within territories of the Usambara Thrush were not associated with habitat structure. We believe the most likely explanation for the Usambara Thrush’s preference for open understorey and closed overstorey habitat relates to foraging behavior. Using information-theoretic model selection we found that invertebrate abundance was the highest-ranked predictor of territory size and was inversely related, consistent with food value theory of territoriality.

Ostrich

Salamander occupancy in headwater stream networks

1. Stream ecosystems exhibit a highly consistent dendritic geometry in which linear habitat units intersect to create a hierarchical network of connected branches. 2. Ecological and life history traits of species living in streams, such as the potential for overland movement, may interact with this architecture to shape patterns of occupancy and response to disturbance. Specifically, large-scale habitat alteration that fragments stream networks and reduces connectivity may reduce the probability a stream is occupied by sensitive species, such as stream salamanders. 3. We collected habitat occupancy data on four species of stream salamanders in first-order (i.e. headwater) streams in undeveloped and urbanised regions of the eastern U.S.A. We then used an information-theoretic approach to test alternative models of salamander occupancy based on a priori predictions of the effects of network configuration, region and salamander life history. 4. Across all four species, we found that streams connected to other first-order streams had higher occupancy than those flowing directly into larger streams and rivers. For three of the four species, occupancy was lower in the urbanised region than in the undeveloped region. 5. These results demonstrate that the spatial configuration of stream networks within protected areas affects the occurrences of stream salamander species. We strongly encourage preservation of network connections between first-order streams in conservation planning and management decisions that may affect stream species.

Freshwater Biology

Linking community-climate disequilibrium to ecosystem function

Turnover in species composition often lags behind the pace of climate change, resulting in mismatches between climate and communities. However, the impact of these community-climate disequilibria on ecosystem functions is rarely considered, and current methods for measuring disequilibria assume that species ranges were, until recently, in equilibrium with climate. Here, we develop a simple theoretical model to address both of these problems by linking community-climate disequilibrium with ecosystem functioning. We show how disequilibrium can impair functioning in the near-term even when climate change is expected to enhance functioning in the long-term. Responses are most likely to change over time in communities where turnover is slow, the impact of disequilibrium counteracts the direct effects of climate on ecosystem function, and pre-existing disequilibrium is large. These findings emphasise the importance of precise and unbiased estimates of community-climate disequilibria for improving ecological forecasts. By fitting our model to time series of both climate and ecosystem function from a metacommunity simulation, we show the potential for community-climate disequilibrium to be inferred without direct knowledge about species' distributions or climatic tolerances. We end by outlining a research agenda to apply dynamic disequilibrium concepts and test novel hypotheses across diverse ecosystems.

Ecology Letters

Environmental correlates of upstream migration of yellow-phase American eels in the Potomac River drainage

Assessing the relationships between upstream migration and environmental variables is important to understanding the ecology of yellow-phase American Eels Anguilla rostrata . During an American Eel migration study within the lower Shenandoah River (Potomac River drainage), we counted and measured American Eels at the Millville Dam eel ladder for three periods: 14 May–23 July 2004, 7–30 September 2004, and 1 June–31 July 2005. Using generalized estimating equations, we modeled each time series of daily American Eel counts by fitting time-varying environmental covariates of lunar illumination (LI), river discharge (RD), and water temperature (WT), including 1-d and 2-d lags of each covariate. Information-theoretic approaches were used for model selection and inference. A total of 4,847 American Eels (19–74 cm total length) used the ladder during the three periods, including 2,622 individuals during a 2-d span following a hurricane-induced peak in river discharge. Additive-effects models of RD + WT, a 2-d lag of LI + RD, and LI + RD were supported for the three periods, respectively. Parameter estimates were positive for river discharge for each time period, negative for lunar illumination for two periods and positive for water temperature during one period. Additive-effects models supported synergistic influences of environmental variables on the upstream migration of yellow-phase American Eels, although river discharge was consistently supported as an influential correlate of upstream migration.

Maryland

Diurnal variations in metal concentrations in the Alamosa River and Wightman Fork, southwestern Colorado, 1995-97

A comprehensive sampling network was implemented in the Alamosa River Basin from 1995 to 1997 to address data gaps identified as part of the ecological risk assessment of the Summitville Superfund site. Aluminum, copper, iron, and zinc were identified as the constituents of concern for the risk assessment. Water-quality samples were collected at six sites on the Alamosa River and Wightman Fork by automatic samplers. Several discrete (instantaneous) samples were collected over 24 hours at each site during periods of high diurnal variations in streamflow (May through September). The discrete samples were analyzed individually and duplicate samples were composited to produce a single sample that represented the daily-mean concentration. The diurnal variations in concentration with respect to the theoretical daily-mean concentration (maximum minus minimum divided by daily mean) are presented. Diurnal metal concentrations were highly variable in the Alamosa River and Wightman Fork. The concentration of a metal at a single site could change by several hundred percent during one diurnal cycle. The largest percent change in metal concentrations was observed for aluminum and iron. Zinc concentrations varied the least of the four metals. No discernible or predictable pattern was indicated in the timing of the daily mean, maximum, or minimum concentrations. The percentage of discrete sample concentrations that varied from the daily-mean concentration by thresholds of plus or minus 10, 25, and 50 percent was evaluated. Between 50 and 75 percent of discrete-sample concentrations varied from the daily-mean concentration by more than plus or minus 10 percent. The percentage of samples exceeding given thresholds generally was smaller during the summer period than the snowmelt period. Sampling strategies are critical to accurately define variability in constituent concentration, and conversely, understanding constituent variability is important in determining appropriate sampling strategies. During nonsteady-state periods, considerable errors in estimates of daily-mean concentration are possible if based on one discrete sample. Flow-weighting multiple discrete samples collected over a diurnal cycle provides a better estimate of daily-mean concentrations during nonsteady-state periods.

Colorado

Sexual selection affects local extinction and turnover in bird communities

Predicting extinction risks has become a central goal for conservation and evolutionary biologists interested in population and community dynamics. Several factors have been put forward to explain risks of extinction, including ecological and life history characteristics of individuals. For instance, factors that affect the balance between natality and mortality can have profound effects on population persistence. Sexual selection has been identified as one such factor. Populations under strong sexual selection experience a number of costs ranging from increased predation and parasitism to enhanced sensitivity to environmental and demographic stochasticity. These findings have led to the prediction that local extinction rates should be higher for species/populations with intense sexual selection. We tested this prediction by analyzing the dynamics of natural bird communities at a continental scale over a period of 21 years (1975-1996), using relevant statistical tools. In agreement with the theoretical prediction, we found that sexual selection increased risks of local extinction (dichromatic birds had on average a 23% higher local extinction rate than monochromatic species). However, despite higher local extinction probabilities, the number of dichromatic species did not decrease over the period considered in this study. This pattern was caused by higher local turnover rates of dichromatic species, resulting in relatively stable communities for both groups of species. Our results suggest that these communities function as metacommunities, with frequent local extinctions followed by colonization. Anthropogenic factors impeding dispersal might therefore have a significant impact on the global persistence of sexually selected species.

Proceedings of the National Academy of Sciences

Fort Collins Science Center- Policy Analysis and Science Assistance Branch : Integrating social, behavioral, economic and biological sciences

The Fort Collins Science Center's Policy Analysis and Science Assistance (PASA) Branch is a team of approximately 22 scientists, technicians, and graduate student researchers. PASA provides unique capabilities in the U.S. Geological Survey by leading projects that integrate social, behavioral, economic, and biological analyses in the context of human-natural resource interactions. Resource planners, managers, and policymakers in the U.S. Departments of the Interior (DOI) and Agriculture (USDA), State and local agencies, as well as international agencies use information from PASA studies to make informed natural resource management and policy decisions. PASA scientists' primary functions are to conduct both theoretical and applied social science research, provide technical assistance, and offer training to advance performance in policy relevant research areas. Management and research issues associated with human-resource interactions typically occur in a unique context, involve difficult to access populations, require knowledge of both natural/biological science in addition to social science, and require the skill to integrate multiple science disciplines. In response to these difficult contexts, PASA researchers apply traditional and state-of-the-art social science methods drawing from the fields of sociology, demography, economics, political science, communications, social-psychology, and applied industrial organization psychology. Social science methods work in concert with our rangeland/agricultural management, wildlife, ecology, and biology capabilities. The goal of PASA's research is to enhance natural resource management, agency functions, policies, and decision-making. Our research is organized into four broad areas of study.

Fact Sheet

Occam's shadow: levels of analysis in evolutionary ecology - where to next?

Evolutionary ecology is the study of evolutionary processes, and the ecological conditions that influence them. A fundamental paradigm underlying the study of evolution is natural selection. Although there are a variety of operational definitions for natural selection in the literature, perhaps the most general one is that which characterizes selection as the process whereby heritable variation in fitness associated with variation in one or more phenotypic traits leads to intergenerational change in the frequency distribution of those traits. The past 20 years have witnessed a marked increase in the precision and reliability of our ability to estimate one or more components of fitness and characterize natural selection in wild populations, owing particularly to significant advances in methods for analysis of data from marked individuals. In this paper, we focus on several issues that we believe are important considerations for the application and development of these methods in the context of addressing questions in evolutionary ecology. First, our traditional approach to estimation often rests upon analysis of aggregates of individuals, which in the wild may reflect increasingly non-random (selected) samples with respect to the trait(s) of interest. In some cases, analysis at the aggregate level, rather than the individual level, may obscure important patterns. While there are a growing number of analytical tools available to estimate parameters at the individual level, and which can cope (to varying degrees) with progressive selection of the sample, the advent of new methods does not reduce the need to consider carefully the appropriate level of analysis in the first place. Estimation should be motivated a priori by strong theoretical analysis. Doing so provides clear guidance, in terms of both (i) assisting in the identification of realistic and meaningful models to include in the candidate model set, and (ii) providing the appropriate context under which the results are interpreted. Second, while it is true that selection (as defined) operates at the level of the individual, the selection gradient is often (if not generally) conditional on the abundance of the population. As such, it may be important to consider estimating transition rates conditional on both the parameter values of the other individuals in the population (or at least their distribution), and population abundance. This will undoubtedly pose a considerable challenge, for both single- and multi-strata applications. It will also require renewed consideration of the estimation of abundance, especially for open populations. Thirdly, selection typically operates on dynamic, individually varying traits. Such estimation may require characterizing fitness in terms of individual plasticity in one or more state variables, constituting analysis of the norms of reaction of individuals to variable environments. This can be quite complex, especially for traits that are under facultative control. Recent work has indicated that the pattern of selection on such traits is conditional on the relative rates of movement among and frequency of spatially heterogeneous habitats, suggesting analyses of evolution of life histories in open populations can be misleading in some cases.

Journal of Applied Statistics

Hydrology: The interdisciplinary science of water

We live in a world where biophysical and social processes are tightly coupled. Hydrologic systems change in response to a variety of natural and human forces such as climate variability and change, water use and water infrastructure, and land cover change. In turn, changes in hydrologic systems impact socioeconomic, ecological, and climate systems at a number of scales, leading to a coevolution of these interlinked systems. The Harvard Water Program, Hydrosociology, Integrated Water Resources Management, Ecohydrology, Hydromorphology, and Sociohydrology were all introduced to provide distinct, interdisciplinary perspectives on water problems to address the contemporary dynamics of human interaction with the hydrosphere and the evolution of the Earth’s hydrologic systems. Each of them addresses scientific, social, and engineering challenges related to how humans influence water systems and vice versa. There are now numerous examples in the literature of how holistic approaches can provide a structure and vision of the future of hydrology. We review selected examples, which taken together, describe the type of theoretical and applied integrated hydrologic analyses and associated curricular content required to address the societal issue of water resources sustainability. We describe a modern interdisciplinary science of hydrology needed to develop an in-depth understanding of the dynamics of the connectedness between human and natural systems and to determine effective solutions to resolve the complex water problems that the world faces today. Nearly, every theoretical hydrologic model introduced previously is in need of revision to accommodate how climate, land, vegetation, and socioeconomic factors interact, change, and evolve over time.

Water Resources Research

A general modeling framework for describing spatially structured population dynamics

Variation in movement across time and space fundamentally shapes the abundance and distribution of populations. Although a variety of approaches model structured population dynamics, they are limited to specific types of spatially structured populations and lack a unifying framework. Here, we propose a unified network-based framework sufficiently novel in its flexibility to capture a wide variety of spatiotemporal processes including metapopulations and a range of migratory patterns. It can accommodate different kinds of age structures, forms of population growth, dispersal, nomadism and migration, and alternative life-history strategies. Our objective was to link three general elements common to all spatially structured populations (space, time and movement) under a single mathematical framework. To do this, we adopt a network modeling approach. The spatial structure of a population is represented by a weighted and directed network. Each node and each edge has a set of attributes which vary through time. The dynamics of our network-based population is modeled with discrete time steps. Using both theoretical and real-world examples, we show how common elements recur across species with disparate movement strategies and how they can be combined under a unified mathematical framework. We illustrate how metapopulations, various migratory patterns, and nomadism can be represented with this modeling approach. We also apply our network-based framework to four organisms spanning a wide range of life histories, movement patterns, and carrying capacities. General computer code to implement our framework is provided, which can be applied to almost any spatially structured population. This framework contributes to our theoretical understanding of population dynamics and has practical management applications, including understanding the impact of perturbations on population size, distribution, and movement patterns. By working within a common framework, there is less chance that comparative analyses are colored by model details rather than general principles

Ecology and Evolution

Evaluating model structure adequacy: The case of the Maggia Valley groundwater system, southern Switzerland

Model adequacy is evaluated with alternative models rated using model selection criteria (AICc, BIC, and KIC) and three other statistics. Model selection criteria are tested with cross-validation experiments and insights for using alternative models to evaluate model structural adequacy are provided. The study is conducted using the computer codes UCODE_2005 and MMA (MultiModel Analysis). One recharge alternative is simulated using the TOPKAPI hydrological model. The predictions evaluated include eight heads and three flows located where ecological consequences and model precision are of concern. Cross-validation is used to obtain measures of prediction accuracy. Sixty-four models were designed deterministically and differ in representation of river, recharge, bedrock topography, and hydraulic conductivity. Results include: (1) What may seem like inconsequential choices in model construction may be important to predictions. Analysis of predictions from alternative models is advised. (2) None of the model selection criteria consistently identified models with more accurate predictions. This is a disturbing result that suggests to reconsider the utility of model selection criteria, and/or the cross-validation measures used in this work to measure model accuracy. (3) KIC displayed poor performance for the present regression problems; theoretical considerations suggest that difficulties are associated with wide variations in the sensitivity term of KIC resulting from the models being nonlinear and the problems being ill-posed due to parameter correlations and insensitivity. The other criteria performed somewhat better, and similarly to each other. (4) Quantities with high leverage are more difficult to predict. The results are expected to be generally applicable to models of environmental systems.

Water Resources Research

Characteristics of foraging sites and protein status in wintering muskoxen: insights from isotopes of nitrogen

Identifying links between nutritional condition of individuals and population trajectories greatly enhances our understanding of the ecology, conservation, and management of wildlife. For northern ungulates, the potential impacts of a changing climate to populations are predicted to be nutritionally mediated through an increase in the severity and variance in winter conditions. Foraging conditions and the availability of body protein as a store for reproduction in late winter may constrain productivity in northern ungulates, yet the link between characteristics of wintering habitats and protein status has not been established for a wild ungulate. We used a non‐invasive proxy of protein status derived from isotopes of N in excreta to evaluate the influence of winter habitats on the protein status of muskoxen in three populations in Alaska (2005–2008). Multiple regression and an information‐theoretic approach were used to compare models that evaluated the influence of population, year, and characteristics of foraging sites (components of diet and physiography) on protein status for groups of muskoxen. The observed variance in protein status among groups of muskoxen across populations and years was partially explained (45%) by local foraging conditions that affected forage availability. Protein status improved for groups of muskoxen as the amount of graminoids in the diet increased (−0.430 ± 0.31, β± 95% CI) and elevation of foraging sites decreased (0.824 ± 0.67). Resources available for reproduction in muskoxen are highly dependent upon demographic, environmental, and physiographic constraints that affect forage availability in winter. Due to their very sedentary nature in winter, muskoxen are highly susceptible to localized foraging conditions; therefore, the spatial variance in resource availability may exert a strong effect on productivity. Consequently, there is a clear need to account for climate–topography effects in winter at multiple scales when predicting the potential impacts of climatic shifts on population trajectories of muskoxen.

Oikos

A test of the Niche Variation Hypothesis in a ruminant herbivore

Despite the shared prediction that the width of a population's dietary niche expands as food becomes limiting, the Niche Variation Hypothesis (NVH) and Optimal Foraging Theory (OFT) offer contrasting views about how individuals alter diet selection when food is limited. Classical OFT predicts that dietary preferences do not change as food becomes limiting, so individuals expand their diets as they compensate for a lack of preferred foods. In contrast, the NVH predicts that among-individual variation in cognition, physiology or morphology create functional trade-offs in foraging efficiency, thereby causing individuals to specialize on different subsets of food as food becomes limiting. To evaluate (a) the predictions of the NVH and OFT and (b) evidence for physiological and cognitive-based functional trade-offs, we used DNA microsatellites and metabarcoding to quantify the diet, microbiome and genetic relatedness (a proxy for social learning) of 218 moose Alces alces across six populations that varied in their degree of food limitation. Consistent with both the NVH and OFT, dietary niche breadth increased with food limitation. Increased diet breadth of individuals—rather than increased diet specialization—was strongly correlated with both food limitation and dietary niche breadth of populations, indicating that moose foraged in accordance with OFT. Diets were not constrained by inheritance of the microbiome or inheritance of diet selection, offering support for the little-tested hypothesis that functional trade-offs in food use (or lack thereof) determine whether populations adhere to the predictions of the NVH or OFT. Our results indicate that both the absence of strong functional trade-offs and the digestive physiology of ruminants provide contexts under which populations should forage in accordance with OFT rather than the NVH. Also, because dietary niche width increased with increased food limitation, OFT and the NVH provide theoretical support for the notion that plant–herbivore interaction networks are plastic rather than static, which has important implications for understanding interspecific niche partitioning. Lastly, because population-level dietary niche breadth and calf recruitment are correlated, and because calf recruitment can be a proxy for food limitation, our work demonstrates how diet data can be employed to understand a populations' proximity to carrying capacity.

Colorado, Utah, Wyoming

Comparisons of patch-use models for wintering American tree sparrows

Optimal foraging theory has stimulated numerous theoretical and empirical studies of foraging behavior for >20 years. These models provide a valuable tool for studying the foraging behavior of an organism. As with any other tool, the models are most effective when properly used. For example, to obtain a robust test of a foraging model, Stephens and Krebs (1986) recommend experimental designs in which four questions are answered in the affirmative. First, do the foragers play the same "game" as the model? Sec- ond, are the assumptions of the model met? Third, does the test rule out alternative possibilities? Finally, are the appropriate variables measured? Negative an- swers to any of these questions could invalidate the model and lead to confusion over the usefulness of foraging theory in conducting ecological studies. Gaines (1989) attempted to determine whether American Tree Sparrows ( Spizella arborea ) foraged by a time (Krebs 1973) or number expectation rule (Gibb 1962), or in a manner consistent with the predictions of Charnov's (1976) marginal value theorem (MVT). Gaines (1989: 118) noted appropriately that field tests of foraging models frequently involve uncontrollable circumstances; thus, it is often difficult to meet the assumptions of the models. Gaines also states (1989: 118) that "violations of the assumptions are also in- formative but do not constitute robust tests of predicted hypotheses," and that "the problem can be avoided by experimental analyses which concurrently test mutually exclusive hypotheses so that alter- native predictions will be eliminated if falsified." There is a problem with this approach because, when major assumptions of models are not satisfied, it is not justifiable to compare a predator's foraging behavior with the model's predictions. I submit that failing to follow the advice offered by Stephens and Krebs (1986) can invalidate tests of foraging models.

The Auk