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The accuracy of ecological flow metrics derived using a physics-based distributed rainfall-runoff model in the Great Plains, USA

The development of a hydrologic foundation, essential for advancing our understanding of flow-ecology relationships, was developed using the high-resolution physics-based distributed rainfall–runoff model V flo in a semi-arid region. We compared the accuracy and bias associated with flow metrics that were generated using V flo , gauge data, and drainage area ratios at both a daily and monthly time step in the Canadian River basin, USA. First, we calibrated and applied bias correction to the V flo model to simulate streamflow at ungauged catchment locations. Next, flow metrics were calculated using simulated and observed data from stream gauge locations. We found discharge predictions using V flo were more accurate than drainage area ratios. General correspondence between predicted discharge and the gauge data was apparent; however, flow metrics calculated using the V flo output did not accurately represent flow variability. Results from the V flo model showed systematic discharge over-predictions in the upper basin and isolated over-predictions in the lower basin, likely due to hail events and sparse rainfall data across the large catchment. Goodness-of-fit statistics (Nash–Sutcliffe efficiency, root-mean square error, and the coefficient of variation) indicated the drainage area ratio and V flo were more accurate at a monthly rather than daily time step, even after quantile mapping. This finding limits the number of streamflow metrics available to develop ecological models, but more importantly, the coarser resolution may hinder our understanding of ecological processes that occur at a submonthly time step. Our approach provides a framework for selecting flow metrics that best represent hydrologic patterns across a large semi-arid catchment with the necessary accuracy to address the ecological questions of interest.

New Mexico, Oklahoma, Texas

Modeling avian abundance from replicated counts using binomial mixture models

Abundance estimation in ecology is usually accomplished by capture–recapture, removal, or distance sampling methods. These may be hard to implement at large spatial scales. In contrast, binomial mixture models enable abundance estimation without individual identification, based simply on temporally and spatially replicated counts. Here, we evaluate mixture models using data from the national breeding bird monitoring program in Switzerland, where some 250 1-km 2 quadrats are surveyed using the territory mapping method three times during each breeding season. We chose eight species with contrasting distribution (wide–narrow), abundance (high–low), and detectability (easy–difficult). Abundance was modeled as a random effect with a Poisson or negative binomial distribution, with mean affected by forest cover, elevation, and route length. Detectability was a logit-linear function of survey date, survey date-by-elevation, and sampling effort (time per transect unit). Resulting covariate effects and parameter estimates were consistent with expectations. Detectability per territory (for three surveys) ranged from 0.66 to 0.94 (mean 0.84) for easy species, and from 0.16 to 0.83 (mean 0.53) for difficult species, depended on survey effort for two easy and all four difficult species, and changed seasonally for three easy and three difficult species. Abundance was positively related to route length in three high-abundance and one low-abundance (one easy and three difficult) species, and increased with forest cover in five forest species, decreased for two nonforest species, and was unaffected for a generalist species. Abundance estimates under the most parsimonious mixture models were between 1.1 and 8.9 (median 1.8) times greater than estimates based on territory mapping; hence, three surveys were insufficient to detect all territories for each species. We conclude that binomial mixture models are an important new approach for estimating abundance corrected for detectability when only repeated-count data are available. Future developments envisioned include estimation of trend, occupancy, and total regional abundance.

Ecological Applications

Assessing tiger population dynamics using photographic capture-recapture sampling

Although wide-ranging, elusive, large carnivore species, such as the tiger, are of scientific and conservation interest, rigorous inferences about their population dynamics are scarce because of methodological problems of sampling populations at the required spatial and temporal scales. We report the application of a rigorous, noninvasive method for assessing tiger population dynamics to test model-based predictions about population viability. We obtained photographic capture histories for 74 individual tigers during a nine-year study involving 5725 trap-nights of effort. These data were modeled under a likelihood-based, ?robust design? capture?recapture analytic framework. We explicitly modeled and estimated ecological parameters such as time-specific abundance, density, survival, recruitment, temporary emigration, and transience, using models that incorporated effects of factors such as individual heterogeneity, trap-response, and time on probabilities of photo-capturing tigers. The model estimated a random temporary emigration parameter of =K' =Y' 0.10 ? 0.069 (values are estimated mean ? SE). When scaled to an annual basis, tiger survival rates were estimated at S = 0.77 ? 0.051, and the estimated probability that a newly caught animal was a transient was = 0.18 ? 0.11. During the period when the sampled area was of constant size, the estimated population size Nt varied from 17 ? 1.7 to 31 ? 2.1 tigers, with a geometric mean rate of annual population change estimated as = 1.03 ? 0.020, representing a 3% annual increase. The estimated recruitment of new animals, Bt, varied from 0 ? 3.0 to 14 ? 2.9 tigers. Population density estimates, D, ranged from 7.33 ? 0.8 tigers/100 km2 to 21.73 ? 1.7 tigers/100 km2 during the study. Thus, despite substantial annual losses and temporal variation in recruitment, the tiger density remained at relatively high levels in Nagarahole. Our results are consistent with the hypothesis that protected wild tiger populations can remain healthy despite heavy mortalities because of their inherently high reproductive potential. The ability to model the entire photographic capture history data set and incorporate reduced-parameter models led to estimates of mean annual population change that were sufficiently precise to be useful. This efficient, noninvasive sampling approach can be used to rigorously investigate the population dynamics of tigers and other elusive, rare, wide-ranging animal species in which individuals can be identified from photographs or other means.

Ecology

A generalized model for estimating the energy density of invertebrates

Invertebrate energy density (ED) values are traditionally measured using bomb calorimetry. However, many researchers rely on a few published literature sources to obtain ED values because of time and sampling constraints on measuring ED with bomb calorimetry. Literature values often do not account for spatial or temporal variability associated with invertebrate ED. Thus, these values can be unreliable for use in models and other ecological applications. We evaluated the generality of the relationship between invertebrate ED and proportion of dry-to-wet mass (pDM). We then developed and tested a regression model to predict ED from pDM based on a taxonomically, spatially, and temporally diverse sample of invertebrates representing 28 orders in aquatic (freshwater, estuarine, and marine) and terrestrial (temperate and arid) habitats from 4 continents and 2 oceans. Samples included invertebrates collected in all seasons over the last 19 y. Evaluation of these data revealed a significant relationship between ED and pDM ( r 2 = 0.96, p < 0.0001), where ED (as J/g wet mass) was estimated from pDM as ED = 22,960pDM &minus; 174.2. Model evaluation showed that nearly all (98.8%) of the variability between observed and predicted values for invertebrate ED could be attributed to residual error in the model. Regression of observed on predicted values revealed that the 97.5% joint confidence region included the intercept of 0 (&minus;103.0 &plusmn; 707.9) and slope of 1 (1.01 &plusmn; 0.12). Use of this model requires that only dry and wet mass measurements be obtained, resulting in significant time, sample size, and cost savings compared to traditional bomb calorimetry approaches. This model should prove useful for a wide range of ecological studies because it is unaffected by taxonomic, seasonal, or spatial variability.

Freshwater Science

A laboratory-calibrated model of coho salmon growth with utility for ecological analyses

We conducted a meta-analysis of laboratory- and hatchery-based growth data to estimate broadly applicable parameters of mass- and temperature-dependent growth of juvenile coho salmon ( Oncorhynchus kisutch ). Following studies of other salmonid species, we incorporated the Ratkowsky growth model into an allometric model and fit this model to growth observations from eight studies spanning ten different populations. To account for changes in growth patterns with food availability, we reparameterized the Ratkowsky model to scale several of its parameters relative to ration. The resulting model was robust across a wide range of ration allocations and experimental conditions, accounting for 99% of the variation in final body mass. We fit this model to growth data from coho salmon inhabiting tributaries and constructed ponds in the Klamath Basin by estimating habitat-specific indices of food availability. The model produced evidence that constructed ponds provided higher food availability than natural tributaries. Because of their simplicity (only mass and temperature are required as inputs) and robustness, ration-varying Ratkowsky models have utility as an ecological tool for capturing growth in freshwater fish populations.

Canadian Journal of Fisheries and Aquatic Sciences

Spatial occupancy models for predicting metapopulation dynamics and viability following reintroduction

The reintroduction of a species into its historic range is a critical component of conservation programmes designed to restore extirpated metapopulations. However, many reintroduction efforts fail, and the lack of rigorous monitoring programmes and statistical models have prevented a general understanding of the factors affecting metapopulation viability following reintroduction. Spatially explicit metapopulation theory provides the basis for understanding the dynamics of fragmented populations linked by dispersal, but the theory has rarely been used to guide reintroduction programmes because most spatial metapopulation models require presence&ndash;absence data from every site in the network, and they do not allow for observation error such as imperfect detection. We develop a spatial occupancy model that relaxes these restrictive assumptions and allows for inference about metapopulation extinction risk and connectivity. We demonstrate the utility of the model using six years of data on the Chiricahua leopard frog Lithobates chiricahuensis , a threatened desert-breeding amphibian that was reintroduced to a network of sites in Arizona USA in 2003. Our results indicate that the model can generate precise predictions of extinction risk and produce connectivity maps that can guide conservation efforts following reintroduction. In the case of L. chiricahuensis , many sites were functionally isolated, and 82% of sites were characterized by intermittent water availability and high local extinction probabilities (0&middot;84, 95% CI: 0&middot;64&ndash;0&middot;99). However, under the current hydrological conditions and spatial arrangement of sites, the risk of metapopulation extinction is estimated to be <3% over a 50-year time horizon. Low metapopulation extinction risk appears to result from the high dispersal capability of the species, the high density of sites in the region and the existence of predator-free permanent wetlands with low local extinction probabilities. Should management be required, extinction risk can be reduced by either increasing the hydroperiod of existing sites or by creating new sites to increase connectivity. Synthesis and applications . This work demonstrates how spatio-temporal statistical models based on ecological theory can be applied to forecast the outcomes of conservation actions such as reintroduction. Our spatial occupancy model should be particularly useful when management agencies lack the funds to collect intensive individual-level data.

Arizona, New Mexico

Creating multithemed ecological regions for macroscale ecology: Testing a flexible, repeatable, and accessible clustering method

Understanding broad-scale ecological patterns and processes often involves accounting for regional-scale heterogeneity. A common way to do so is to include ecological regions in sampling schemes and empirical models. However, most existing ecological regions were developed for specific purposes, using a limited set of geospatial features and irreproducible methods. Our study purpose was to: (1) describe a method that takes advantage of recent computational advances and increased availability of regional and global data sets to create customizable and reproducible ecological regions, (2) make this algorithm available for use and modification by others studying different ecosystems, variables of interest, study extents, and macroscale ecology research questions, and (3) demonstrate the power of this approach for the research question—How well do these regions capture regional-scale variation in lake water quality? To achieve our purpose we: (1) used a spatially constrained spectral clustering algorithm that balances geospatial homogeneity and region contiguity to create ecological regions using multiple terrestrial, climatic, and freshwater geospatial data for 17 northeastern U.S. states (~1,800,000 km 2 ); (2) identified which of the 52 geospatial features were most influential in creating the resulting 100 regions; and (3) tested the ability of these ecological regions to capture regional variation in water nutrients and clarity for ~6,000 lakes. We found that: (1) a combination of terrestrial, climatic, and freshwater geospatial features influenced region creation, suggesting that the oft-ignored freshwater landscape provides novel information on landscape variability not captured by traditionally used climate and terrestrial metrics; and (2) the delineated regions captured macroscale heterogeneity in ecosystem properties not included in region delineation—approximately 40% of the variation in total phosphorus and water clarity among lakes was at the regional scale. Our results demonstrate the usefulness of this method for creating customizable and reproducible regions for research and management applications.

Ecology and Evolution

Ecosystem effects of environmental flows: Modelling and experimental floods in a dryland river

Successful environmental flow prescriptions require an accurate understanding of the linkages among flow events, geomorphic processes and biotic responses. We describe models and results from experimental flow releases associated with an environmental flow program on the Bill Williams River (BWR), Arizona, in arid to semiarid western U.S.A. Two general approaches for improving knowledge and predictions of ecological responses to environmental flows are: (1) coupling physical system models to ecological responses and (2) clarifying empirical relationships between flow and ecological responses through implementation and monitoring of experimental flow releases. We modelled the BWR physical system using: (1) a reservoir operations model to simulate reservoir releases and reservoir water levels and estimate flow through the river system under a range of scenarios, (2) one- and two-dimensional river hydraulics models to estimate stage-discharge relationships at the whole-river and local scales, respectively, and (3) a groundwater model to estimate surface- and groundwater interactions in a large, alluvial valley on the BWR where surface flow is frequently absent. An example of a coupled, hydrology-ecology model is the Ecosystems Function Model, which we used to link a one-dimensional hydraulic model with riparian tree seedling establishment requirements to produce spatially explicit predictions of seedling recruitment locations in a Geographic Information System. We also quantified the effects of small experimental floods on the differential mortality of native and exotic riparian trees, on beaver dam integrity and distribution, and on the dynamics of differentially flow-adapted benthic macroinvertebrate groups. Results of model applications and experimental flow releases are contributing to adaptive flow management on the BWR and to the development of regional environmental flow standards. General themes that emerged from our work include the importance of response thresholds, which are commonly driven by geomorphic thresholds or mediated by geomorphic processes, and the importance of spatial and temporal variation in the effects of flows on ecosystems, which can result from factors such as longitudinal complexity and ecohydrological feedbacks. ?? Published 2009.

Freshwater Biology

Ecological site-based assessments of wind and water erosion: informing accelerated soil erosion management in rangelands

Accelerated soil erosion occurs when anthropogenic processes modify soil, vegetation or climatic conditions causing erosion rates at a location to exceed their natural variability. Identifying where and when accelerated erosion occurs is a critical first step toward its effective management. Here we explore how erosion assessments structured in the context of ecological sites (a land classification based on soils, landscape setting and ecological potential) and their vegetation states (plant assemblages that may change due to management) can inform systems for reducing accelerated soil erosion in rangelands. We evaluated aeolian horizontal sediment flux and fluvial sediment erosion rates for five ecological sites in southern New Mexico, USA, using monitoring data and rangeland-specific wind and water erosion models. Across the ecological sites, plots in shrub-encroached and shrub-dominated vegetation states were consistently susceptible to aeolian sediment flux and fluvial sediment erosion. Both processes were found to be highly variable for grassland and grass-succulent states across the ecological sites at the plot scale (0.25 Ha). We identify vegetation thresholds that define cover levels below which rapid (exponential) increases in aeolian sediment flux and fluvial sediment erosion occur across the ecological sites and vegetation states. Aeolian sediment flux and fluvial erosion in the study area can be effectively controlled when bare ground cover is <20% of a site or the cover of canopy interspaces >100 cm in length is less than ~35%. Land use and management activities that alter cover levels such that they cross thresholds, and/or drive vegetation state changes, may increase the susceptibility of areas to erosion. Land use impacts that are constrained within the range of natural variability should not result in accelerated soil erosion. Evaluating land condition against the erosion thresholds identified here will enable identification of areas susceptible to accelerated soil erosion and the development of practical management solutions.

New Mexico

Optimal temperature for malaria transmission is dramaticallylower than previously predicted

The ecology of mosquito vectors and malaria parasites affect the incidence, seasonal transmission and geographical range of malaria. Most malaria models to date assume constant or linear responses of mosquito and parasite life-history traits to temperature, predicting optimal transmission at 31 °C. These models are at odds with field observations of transmission dating back nearly a century. We build a model with more realistic ecological assumptions about the thermal physiology of insects. Our model, which includes empirically derived nonlinear thermal responses, predicts optimal malaria transmission at 25 °C (6 °C lower than previous models). Moreover, the model predicts that transmission decreases dramatically at temperatures > 28 °C, altering predictions about how climate change will affect malaria. A large data set on malaria transmission risk in Africa validates both the 25 °C optimum and the decline above 28 °C. Using these more accurate nonlinear thermal-response models will aid in understanding the effects of current and future temperature regimes on disease transmission.

Ecology Letters

Hydrodynamic model of the Colorado River, Glen Canyon Dam to Lees Ferry in Glen Canyon National Recreation Area, Arizona

The U.S. Geological Survey constructed a two-dimensional hydrodynamic model that was applied to a 15.8-mile tailwater reach of the Colorado River in Glen Canyon that begins 0.25 mile downstream from Glen Canyon Dam and extends to Lees Ferry in Glen Canyon National Recreation Area, Arizona. The model used the Flow and Sediment Transport with Morphological Evolution of Channels (FaSTMECH) solver in the International River Interface Cooperative (iRIC) modeling interface. The model grid was developed from a full channel digital elevation model derived by combining bathymetric and topographic data collected from March 2013 to February 2016. The model was used to predict water-surface elevations, depths, depth-averaged flow velocities, and bed shear stresses for discharges ranging from 1,000 to 70,000 cubic feet per second. Modeled water-surface elevations matched well with measured values at cross sections throughout the reach, with a mean absolute error of 0.14 meter over the range of typical discharge releases from Glen Canyon Dam. The mean error on discharge, a measure of how well the model solution converged, averaged 0.6 percent and did not exceed 2 percent over the range of discharges modeled. These results indicate that model predictions of hydraulic parameters are reasonably accurate and suitable for use for a variety of purposes, such as ecological and geomorphic modeling.

Arizona

Climate patterns as predictors of amphibians species richness and indicators of potential stress

Amphibians occupy a range of habitats throughout the world, but species richness is greatest in regions with moist, warm climates. We modeled the statistical relations of anuran and urodele species richness with mean annual climate for the conterminous United States, and compared the strength of these relations at national and regional levels. Model variables were calculated for county and subcounty mapping units, and included 40-year (1960-1999) annual mean and mean annual climate statistics, mapping unit average elevation, mapping unit land area, and estimates of anuran and urodele species richness. Climate data were derived from more than 7,500 first-order and cooperative meteorological stations and were interpolated to the mapping units using multiple linear regression models. Anuran and urodele species richness were calculated from the United States Geological Survey's Amphibian Research and Monitoring Initiative (ARMI) National Atlas for Amphibian Distributions. The national multivariate linear regression (MLR) model of anuran species richness had an adjusted coefficient of determination (R2) value of 0.64 and the national MLR model for urodele species richness had an R2 value of 0.45. Stratifying the United States by coarse-resolution ecological regions provided models for anUrans that ranged in R2 values from 0.15 to 0.78. Regional models for urodeles had R2 values. ranging from 0.27 to 0.74. In general, regional models for anurans were more strongly influenced by temperature variables, whereas precipitation variables had a larger influence on urodele models.

Alytes

Ecological effects on streams from forest fertilization: Literature review and conceptual framework for future study in the western Cascades

Fertilization of forests with urea-nitrogen has been studied numerous times for its effects on water quality. Stream nitrogen concentrations following fertilization are typically elevated during winter, including peaks in the tens-of-thousands of parts per billion range, with summer concentrations often returning to background or near-background levels. Despite these increases, water-quality criteria for nitrogen have rarely been exceeded. However, such criteria are targeted at fish toxicity or human health and are not relevant to concentrations that could cause ecological disturbances. Studies of the responses of stream biota to fertilization have been rare and have targeted either immediate, toxicity-based responses or used methods insensitive to ongoing ecological processes. This report reviews water-quality studies following forest fertilizations, emphasizing Cascade streams in the Pacific Northwest and documented biological responses in those streams. A conceptual model predicting potential ecological response to fertilization, which includes effects on algal growth and primary production, is presented. In this model, applied fertilizer nitrogen reaching streams is mostly exported during winter. However, some nitrogen retained in soils or stream and riparian areas may become available to aquatic biota during spring and summer. Biological responses may be minimal in small streams nearest to application because of light limitation, but may be elevated downstream where light is sufficient to allow algal growth. Ultimately, algal response could be greatest in downstream reaches, although ambient nutrient concentrations remain low due to uptake and benthic nutrient recycling. Ground-water flow paths and hyporheic processing could be critical in determining the fate of applied nitrogen. A framework is provided for testing this response in the Little River watershed, a tributary to the North Umpqua River, Oregon, at basic and intensive levels of investigation.

Oregon

The rise of an apex predator following deglaciation

Aim Sea otters ( Enhydra lutris ) are an apex predator of the nearshore marine community and nearly went extinct at the turn of the 20th century. Reintroductions and legal protection allowed sea otters to re‐colonize much of their former range. Our objective was to chronicle the colonization of this apex predator in Glacier Bay, Alaska, to help understand the mechanisms that governed their successful colonization. Location Glacier Bay is a tidewater glacier fjord in southeastern Alaska that was entirely covered by glaciers in the mid‐18th century. Since then, it has endured the fastest tidewater glacier retreat in recorded history. Methods We collected and analysed several data sets, spanning 20 years, to document the spatio‐temporal dynamics of an apex predator expanding into an area where they were formerly absent. We used novel quantitative tools to model the occupancy, abundance and colonization dynamics of sea otters, while accounting for uncertainty in the data collection process, the ecological process and model parameters. Results Twenty years after sea otters were first observed colonizing Glacier Bay, they became one of the most abundant and widely distributed marine mammal. The population grew exponentially at a rate of 20% per year. They colonized Glacier Bay at a maximum rate of 6 km per year, with faster colonization rates occurring early in the colonization process. During colonization, sea otters selected shallow areas, close to shore, with a steep bottom slope, and a relatively simple shoreline complexity index. Main conclusions The growth and expansion of sea otters in Glacier Bay demonstrate how legal protection and translocation of apex predators can facilitate their successful establishment into a community in which they were formerly absent. The success of sea otters was, in part, a consequence of habitat that was left largely unperturbed by humans for the past 250 years. Further, sea otters and other marine predators, whose distribution is limited by ice, have the potential to expand in distribution and abundance, reshaping future marine communities in the wake of deglaciation and global loss of sea ice.

Alaska

Linking occurrence and fitness to persistence: Habitat-based approach for endangered Greater Sage-Grouse

Detailed empirical models predicting both species occurrence and fitness across a landscape are necessary to understand processes related to population persistence. Failure to consider both occurrence and fitness may result in incorrect assessments of habitat importance leading to inappropriate management strategies. We took a two-stage approach to identifying critical nesting and brood-rearing habitat for the endangered Greater Sage-Grouse (Centrocercus urophasianus) in Alberta at a landscape scale. First, we used logistic regression to develop spatial models predicting the relative probability of use (occurrence) for Sage-Grouse nests and broods. Secondly, we used Cox proportional hazards survival models to identify the most risky habitats across the landscape. We combined these two approaches to identify Sage-Grouse habitats that pose minimal risk of failure (source habitats) and attractive sink habitats that pose increased risk (ecological traps). Our models showed that Sage-Grouse select for heterogeneous patches of moderate sagebrush cover (quadratic relationship) and avoid anthropogenic edge habitat for nesting. Nests were more successful in heterogeneous habitats, but nest success was independent of anthropogenic features. Similarly, broods selected heterogeneous high-productivity habitats with sagebrush while avoiding human developments, cultivated cropland, and high densities of oil wells. Chick mortalities tended to occur in proximity to oil and gas developments and along riparian habitats. For nests and broods, respectively, approximately 10% and 5% of the study area was considered source habitat, whereas 19% and 15% of habitat was attractive sink habitat. Limited source habitats appear to be the main reason for poor nest success (39%) and low chick survival (12%). Our habitat models identify areas of protection priority and areas that require immediate management attention to enhance recruitment to secure the viability of this population. This novel approach to habitat-based population viability modeling has merit for many species of concern. ?? 2007 by the Ecological Society of America.

Ecological Applications

Supporting dryland restoration success with applied ecological forecasting of seeding outcomes

Introduction Ecological restoration is increasingly used to sustain biodiversity and ecosystem services. In drylands of the western United States (US), post-disturbance restoration often involves seeding treatments to promote the recovery of native plant communities. Spatial and temporal variability in environmental conditions influences plant establishment and contributes to low restoration success in certain locations and years. Objectives Here, we discuss how forecasts for plant establishment can be developed and delivered to help land managers anticipate the impacts of near-term (months to years) environmental conditions on restoration. Methods We developed an ecological forecast system that predicts the outcome of restoration seeding by integrating weather forecasts, an ecosystem water balance model, and plant establishment models. Results In this article, we focus on a conceptual approach to developing, delivering, and applying ecological forecasts for restoration. We illustrate the potential of this approach by adapting existing ecological models to build an initial version of a decision support tool that delivers a species-specific ecological forecast for big sagebrush ( Artemisia tridentata ) establishment. Integrating ecological forecasts into plans for restoration seeding presents opportunities to anticipate and account for environmental variability. Conclusions Finally, we discuss how connecting research, forecast delivery, and management can maximize the impact of ecological forecasts on restoration success.

Idaho, Nevada

Using structural equation modeling to link human activities to wetland ecological integrity

The integrity of wetlands is of global concern. A common approach to evaluating ecological integrity involves bioassessment procedures that quantify the degree to which communities deviate from historical norms. While helpful, bioassessment provides little information about how altered conditions connect to community response. More detailed information is needed for conservation and restoration. We have illustrated an approach to addressing this challenge using structural equation modeling (SEM) and long-term monitoring data from Rocky Mountain National Park (RMNP). Wetlands in RMNP are threatened by a complex history of anthropogenic disturbance including direct alteration of hydrologic regimes; elimination of elk, wolves, and grizzly bears; reintroduction of elk (absent their primary predators); and the extirpation of beaver. More recently, nonnative moose were introduced to the region and have expanded into the park. Bioassessment suggests that up to half of the park's wetlands are not in reference condition. We developed and evaluated a general hypothesis about how human alterations influence wetland integrity and then develop a specific model using RMNP wetlands. Bioassessment revealed three bioindicators that appear to be highly sensitive to human disturbance (HD): (1) conservatism, (2) degree of invasion, and (3) cover of native forbs. SEM analyses suggest several ways human activities have impacted wetland integrity and the landscape of RMNP. First, degradation is highest where the combined effects of all types of direct HD have been the greatest (i.e., there is a general, overall effect). Second, specific HDs appear to create a “mixed-bag” of complex indirect effects, including reduced invasion and increased conservatism, but also reduced native forb cover. Some of these effects are associated with alterations to hydrologic regimes, while others are associated with altered shrub production. Third, landscape features created by historical beaver activity continue to influence wetland integrity years after beavers have abandoned sites via persistent landforms and reduced biomass of tall shrubs. Our model provides a system-level perspective on wetland integrity and provides a context for future evaluations and investigations. It also suggests scientifically supported natural resource management strategies that can assist in the National Park Service mission of maintaining or, when indicated, restoring ecological integrity “unimpaired for future generations.”

Colorado

Trait-based approaches in the analysis of stream fish communities

Species traits are used to study the functional organization of fish communities for a range of reasons, from simply reducing data dimensionality to providing mechanistic explanations for observed variation in communities. Ecological and life history traits have been used to understand the basic ecology of fishes and predict (1) species and community responses to habitat and climate alteration, and (2) species extinction, species invasion, and community homogenization. Many approaches in this arena have been developed during the past three decades, but they often have not been integrated with related ecological concepts or subdisciplines, which has led to confusion in terminology. We review 102 studies of species traits and then summarize patterns in traits being used and questions being addressed with trait-based approaches. Overall, studies of fish–habitat relationships that apply habitat templates and hierarchical filters dominate our sample; the most frequently used traits are related to feeding. We define and show the relationships among key terms such as fundamental and realized niches; functional traits, performance, and fitness; tactic, trait-state, syndromes, and strategies; and guilds and functional groups. We propose accelerating research to (1) quantify trait plasticity, (2) identify traits useful for testing ecological hypotheses, (3) model habitat and biotic interactions in communities while explicitly accounting for phylogenetic relationships, (4) explore how traits control community assembly, and (5) document the importance of traits in fish– community responses to anthropogenic change and in delivering ecosystem services. Further synthesis of these topics is still needed to develop concepts, models, and principles that can unify the disparate approaches taken in trait-based analysis of fish communities, link fish community ecology to general community ecology, and inform sustainable management of ecosystems.

Conference Paper