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T.C. Edwards

Publications and source records attributed to T.C. Edwards.

14 recordsLinked to original sources

Sampling in ecology and evolution - bridging the gap between theory and practice

Sampling is a key issue for answering most ecological and evolutionary questions. The importance of developing a rigorous sampling design tailored to specific questions has already been discussed in the ecological and sampling literature and has provided useful tools and recommendations to sample and analyse ecological data. However, sampling issues are often difficult to overcome in ecological studies due to apparent inconsistencies between theory and practice, often leading to the implementation of simplified sampling designs that suffer from unknown biases. Moreover, we believe that classical sampling principles which are based on estimation of means and variances are insufficient to fully address many ecological questions that rely on estimating relationships between a response and a set of predictor variables over time and space. Our objective is thus to highlight the importance of selecting an appropriate sampling space and an appropriate sampling design. We also emphasize the importance of using prior knowledge of the study system to estimate models or complex parameters and thus better understand ecological patterns and processes generating these patterns. Using a semi-virtual simulation study as an illustration we reveal how the selection of the space (e.g. geographic, climatic), in which the sampling is designed, influences the patterns that can be ultimately detected. We also demonstrate the inefficiency of common sampling designs to reveal response curves between ecological variables and climatic gradients. Further, we show that response-surface methodology, which has rarely been used in ecology, is much more efficient than more traditional methods. Finally, we discuss the use of prior knowledge, simulation studies and model-based designs in defining appropriate sampling designs. We conclude by a call for development of methods to unbiasedly estimate nonlinear ecologically relevant parameters, in order to make inferences while fulfilling requirements of both sampling theory and field work logistics. ?? 2010 The Authors.

Ecography

Hierarchical spatial models for predicting pygmy rabbit distribution and relative abundance

Conservationists routinely use species distribution models to plan conservation, restoration and development actions, while ecologists use them to infer process from pattern. These models tend to work well for common or easily observable species, but are of limited utility for rare and cryptic species. This may be because honest accounting of known observation bias and spatial autocorrelation are rarely included, thereby limiting statistical inference of resulting distribution maps. We specified and implemented a spatially explicit Bayesian hierarchical model for a cryptic mammal species (pygmy rabbit Brachylagus idahoensis). Our approach used two levels of indirect sign that are naturally hierarchical (burrows and faecal pellets) to build a model that allows for inference on regression coefficients as well as spatially explicit model parameters. We also produced maps of rabbit distribution (occupied burrows) and relative abundance (number of burrows expected to be occupied by pygmy rabbits). The model demonstrated statistically rigorous spatial prediction by including spatial autocorrelation and measurement uncertainty. We demonstrated flexibility of our modelling framework by depicting probabilistic distribution predictions using different assumptions of pygmy rabbit habitat requirements. Spatial representations of the variance of posterior predictive distributions were obtained to evaluate heterogeneity in model fit across the spatial domain. Leave-one-out cross-validation was conducted to evaluate the overall model fit. Synthesis and applications. Our method draws on the strengths of previous work, thereby bridging and extending two active areas of ecological research: species distribution models and multi-state occupancy modelling. Our framework can be extended to encompass both larger extents and other species for which direct estimation of abundance is difficult. ?? 2010 The Authors. Journal compilation ?? 2010 British Ecological Society.

Journal of Applied Ecology

Climatic extremes improve predictions of spatial patterns of tree species

Understanding niche evolution, dynamics, and the response of species to climate change requires knowledge of the determinants of the environmental niche and species range limits. Mean values of climatic variables are often used in such analyses. In contrast, the increasing frequency of climate extremes suggests the importance of understanding their additional influence on range limits. Here, we assess how measures representing climate extremes (i.e., interannual variability in climate parameters) explain and predict spatial patterns of 11 tree species in Switzerland. We find clear, although comparably small, improvement (+20% in adjusted D 2 , +8% and +3% in cross-validated True Skill Statistic and area under the receiver operating characteristics curve values) in models that use measures of extremes in addition to means. The primary effect of including information on climate extremes is a correction of local overprediction and underprediction. Our results demonstrate that measures of climate extremes are important for understanding the climatic limits of tree species and assessing species niche characteristics. The inclusion of climate variability likely will improve models of species range limits under future conditions, where changes in mean climate and increased variability are expected.

Conference Paper

Remote sensing-based predictors improve distribution models of rare, early successional and broadleaf tree species in Utah

1. Compared to bioclimatic variables, remote sensing predictors are rarely used for predictive species modelling. When used, the predictors represent typically habitat classifications or filters rather than gradual spectral, surface or biophysical properties. Consequently, the full potential of remotely sensed predictors for modelling the spatial distribution of species remains unexplored. Here we analysed the partial contributions of remotely sensed and climatic predictor sets to explain and predict the distribution of 19 tree species in Utah. We also tested how these partial contributions were related to characteristics such as successional types or species traits. 2. We developed two spatial predictor sets of remotely sensed and topo-climatic variables to explain the distribution of tree species. We used variation partitioning techniques applied to generalized linear models to explore the combined and partial predictive powers of the two predictor sets. Non-parametric tests were used to explore the relationships between the partial model contributions of both predictor sets and species characteristics. 3. More than 60% of the variation explained by the models represented contributions by one of the two partial predictor sets alone, with topo-climatic variables outperforming the remotely sensed predictors. However, the partial models derived from only remotely sensed predictors still provided high model accuracies, indicating a significant correlation between climate and remote sensing variables. The overall accuracy of the models was high, but small sample sizes had a strong effect on cross-validated accuracies for rare species. 4. Models of early successional and broadleaf species benefited significantly more from adding remotely sensed predictors than did late seral and needleleaf species. The core-satellite species types differed significantly with respect to overall model accuracies. Models of satellite and urban species, both with low prevalence, benefited more from use of remotely sensed predictors than did the more frequent core species. 5. Synthesis and applications. If carefully prepared, remotely sensed variables are useful additional predictors for the spatial distribution of trees. Major improvements resulted for deciduous, early successional, satellite and rare species. The ability to improve model accuracy for species having markedly different life history strategies is a crucial step for assessing effects of global change. ?? 2007 The Authors.

Journal of Applied Ecology

Habitat classification modeling with incomplete data: Pushing the habitat envelope

Habitat classification models (HCMs) are invaluable tools for species conservation, land-use planning, reserve design, and metapopulation assessments, particularly at broad spatial scales. However, species occurrence data are often lacking and typically limited to presence points at broad scales. This lack of absence data precludes the use of many statistical techniques for HCMs. One option is to generate pseudo-absence points so that the many available statistical modeling tools can be used. Traditional techniques generate pseudoabsence points at random across broadly defined species ranges, often failing to include biological knowledge concerning the species-habitat relationship. We incorporated biological knowledge of the species-habitat relationship into pseudo-absence points by creating habitat envelopes that constrain the region from which points were randomly selected. We define a habitat envelope as an ecological representation of a species, or species feature's (e.g., nest) observed distribution (i.e., realized niche) based on a single attribute, or the spatial intersection of multiple attributes. We created HCMs for Northern Goshawk (Accipiter gentilis atricapillus) nest habitat during the breeding season across Utah forests with extant nest presence points and ecologically based pseudo-absence points using logistic regression. Predictor variables were derived from 30-m USDA Landfire and 250-m Forest Inventory and Analysis (FIA) map products. These habitat-envelope-based models were then compared to null envelope models which use traditional practices for generating pseudo-absences. Models were assessed for fit and predictive capability using metrics such as kappa, thresholdindependent receiver operating characteristic (ROC) plots, adjusted deviance (Dadj2), and cross-validation, and were also assessed for ecological relevance. For all cases, habitat envelope-based models outperformed null envelope models and were more ecologically relevant, suggesting that incorporating biological knowledge into pseudo-absence point generation is a powerful tool for species habitat assessments. Furthermore, given some a priori knowledge of the species-habitat relationship, ecologically based pseudo-absence points can be applied to any species, ecosystem, data resolution, and spatial extent. ?? 2007 by the Ecological Society of America.

Ecological Applications

Random forests for classification in ecology

Classification procedures are some of the most widely used statistical methods in ecology. Random forests (RF) is a new and powerful statistical classifier that is well established in other disciplines but is relatively unknown in ecology. Advantages of RF compared to other statistical classifiers include (1) very high classification accuracy; (2) a novel method of determining variable importance; (3) ability to model complex interactions among predictor variables; (4) flexibility to perform several types of statistical data analysis, including regression, classification, survival analysis, and unsupervised learning; and (5) an algorithm for imputing missing values. We compared the accuracies of RF and four other commonly used statistical classifiers using data on invasive plant species presence in Lava Beds National Monument, California, USA, rare lichen species presence in the Pacific Northwest, USA, and nest sites for cavity nesting birds in the Uinta Mountains, Utah, USA. We observed high classification accuracy in all applications as measured by cross-validation and, in the case of the lichen data, by independent test data, when comparing RF to other common classification methods. We also observed that the variables that RF identified as most important for classifying invasive plant species coincided with expectations based on the literature. ?? 2007 by the Ecological Society of America.

Ecology

Effects of sample survey design on the accuracy of classification tree models in species distribution models

We evaluated the effects of probabilistic (hereafter DESIGN) and non-probabilistic (PURPOSIVE) sample surveys on resultant classification tree models for predicting the presence of four lichen species in the Pacific Northwest, USA. Models derived from both survey forms were assessed using an independent data set (EVALUATION). Measures of accuracy as gauged by resubstitution rates were similar for each lichen species irrespective of the underlying sample survey form. Cross-validation estimates of prediction accuracies were lower than resubstitution accuracies for all species and both design types, and in all cases were closer to the true prediction accuracies based on the EVALUATION data set. We argue that greater emphasis should be placed on calculating and reporting cross-validation accuracy rates rather than simple resubstitution accuracy rates. Evaluation of the DESIGN and PURPOSIVE tree models on the EVALUATION data set shows significantly lower prediction accuracy for the PURPOSIVE tree models relative to the DESIGN models, indicating that non-probabilistic sample surveys may generate models with limited predictive capability. These differences were consistent across all four lichen species, with 11 of the 12 possible species and sample survey type comparisons having significantly lower accuracy rates. Some differences in accuracy were as large as 50%. The classification tree structures also differed considerably both among and within the modelled species, depending on the sample survey form. Overlap in the predictor variables selected by the DESIGN and PURPOSIVE tree models ranged from only 20% to 38%, indicating the classification trees fit the two evaluated survey forms on different sets of predictor variables. The magnitude of these differences in predictor variables throws doubt on ecological interpretation derived from prediction models based on non-probabilistic sample surveys. ?? 2006 Elsevier B.V. All rights reserved.

Ecological Modelling

Predicting tree species presence and basal area in Utah: A comparison of stochastic gradient boosting, generalized additive models, and tree-based methods

Many efforts are underway to produce broad-scale forest attribute maps by modelling forest class and structure variables collected in forest inventories as functions of satellite-based and biophysical information. Typically, variants of classification and regression trees implemented in Rulequest's?? See5 and Cubist (for binary and continuous responses, respectively) are the tools of choice in many of these applications. These tools are widely used in large remote sensing applications, but are not easily interpretable, do not have ties with survey estimation methods, and use proprietary unpublished algorithms. Consequently, three alternative modelling techniques were compared for mapping presence and basal area of 13 species located in the mountain ranges of Utah, USA. The modelling techniques compared included the widely used See5/Cubist, generalized additive models (GAMs), and stochastic gradient boosting (SGB). Model performance was evaluated using independent test data sets. Evaluation criteria for mapping species presence included specificity, sensitivity, Kappa, and area under the curve (AUC). Evaluation criteria for the continuous basal area variables included correlation and relative mean squared error. For predicting species presence (setting thresholds to maximize Kappa), SGB had higher values for the majority of the species for specificity and Kappa, while GAMs had higher values for the majority of the species for sensitivity. In evaluating resultant AUC values, GAM and/or SGB models had significantly better results than the See5 models where significant differences could be detected between models. For nine out of 13 species, basal area prediction results for all modelling techniques were poor (correlations less than 0.5 and relative mean squared errors greater than 0.8), but SGB provided the most stable predictions in these instances. SGB and Cubist performed equally well for modelling basal area for three species with moderate prediction success, while all three modelling tools produced comparably good predictions (correlation of 0.68 and relative mean squared error of 0.56) for one species. ?? 2006 Elsevier B.V. All rights reserved.

Ecological Modelling

A variance-decomposition approach to investigating multiscale habitat associations

The recognition of the importance of spatial scale in ecology has led many researchers to take multiscale approaches to studying habitat associations. However, few of the studies that investigate habitat associations at multiple spatial scales have considered the potential effects of cross-scale correlations in measured habitat variables. When cross-scale correlations in such studies are strong, conclusions drawn about the relative strength of habitat associations at different spatial scales may be inaccurate. Here we adapt and demonstrate an analytical technique based on variance decomposition for quantifying the influence of cross-scale correlations on multiscale habitat associations. We used the technique to quantify the variation in nest-site locations of Red-naped Sapsuckers (Sphyrapicus nuchalis) and Northern Flickers (Colaptes auratus) associated with habitat descriptors at three spatial scales. We demonstrate how the method can be used to identify components of variation that are associated only with factors at a single spatial scale as well as shared components of variation that represent cross-scale correlations. Despite the fact that no explanatory variables in our models were highly correlated (r < 0.60), we found that shared components of variation reflecting cross-scale correlations accounted for roughly half of the deviance explained by the models. These results highlight the importance of both conducting habitat analyses at multiple spatial scales and of quantifying the effects of cross-scale correlations in such analyses. Given the limits of conventional analytical techniques, we recommend alternative methods, such as the variance-decomposition technique demonstrated here, for analyzing habitat associations at multiple spatial scales. ?? The Cooper Ornithological Society 2006.

Conference Paper

Effect of different sampling schemes on the spatial placement of conservation reserves in Utah, USA

We evaluated the effect of three different sampling schemes used to organize spatially explicit biological information had on the spatial placement of conservation reserves in Utah, USA. The three sampling schemes consisted of a hexagon representation developed by the EPA/EMAP program (statistical basis), watershed boundaries (ecological), and the current county boundaries of Utah (socio-political). Four decision criteria were used to estimate effects, including amount of area, length of edge, lowest number of contiguous reserves, and greatest number of terrestrial vertebrate species covered. A fifth evaluation criterion was the effect each sampling scheme had on the ability of the modeled conservation reserves to cover the six major ecoregions found in Utah. Of the three sampling schemes, county boundaries covered the greatest number of species, but also created the longest length of edge and greatest number of reserves. Watersheds maximized species coverage using the least amount of area. Hexagons and watersheds provide the least amount of edge and fewest number of reserves. Although there were differences in area, edge and number of reserves among the sampling schemes, all three schemes covered all the major ecoregions in Utah and their inclusive biodiversity. ?? 2003 Elsevier Science Ltd. All rights reserved.

Biological Conservation

Generalized linear and generalized additive models in studies of species distributions: Setting the scene

An important statistical development of the last 30 years has been the advance in regression analysis provided by generalized linear models (GLMs) and generalized additive models (GAMs). Here we introduce a series of papers prepared within the framework of an international workshop entitled: Advances in GLMs/GAMs modeling: from species distribution to environmental management, held in Riederalp, Switzerland, 6-11 August 2001. We first discuss some general uses of statistical models in ecology, as well as provide a short review of several key examples of the use of GLMs and GAMs in ecological modeling efforts. We next present an overview of GLMs and GAMs, and discuss some of their related statistics used for predictor selection, model diagnostics, and evaluation. Included is a discussion of several new approaches applicable to GLMs and GAMs, such as ridge regression, an alternative to stepwise selection of predictors, and methods for the identification of interactions by a combined use of regression trees and several other approaches. We close with an overview of the papers and how we feel they advance our understanding of their application to ecological modeling. ?? 2002 Elsevier Science B.V. All rights reserved.

Ecological Modelling

Landscape patterns as habitat predictors: Building and testing models for cavity-nesting birds in the Uinta Mountains of Utah, USA

The ability to predict species occurrences quickly is often crucial for managers and conservation biologists with limited time and funds. We used measured associations with landscape patterns to build accurate predictive habitat models that were quickly and easily applied (i.e., required no additional data collection in the field to make predictions). We used classification trees (a nonparametric alternative to discriminant function analysis, logistic regression, and other generalized linear models) to model nesting habitat of red-naped sapsuckers (Sphyrapicus nuchalis), northern flickers (Colaptes auratus), tree swallows (Tachycineta bicolor), and mountain chickadees (Parus gambeli) in the Uinta Mountains of northeastern Utah, USA. We then tested the predictive capability of the models with independent data collected in the field the following year. The models built for the northern flicker, red-naped sapsucker, and tree swallow were relatively accurate (84%, 80%, and 75% nests correctly classified, respectively) compared to the models for the mountain chickadee (50% nests correctly classified). All four models were more selective than a null model that predicted habitat based solely on a gross association with aspen forests. We conclude that associations with landscape patterns can be used to build relatively accurate, easy to use, predictive models for some species. Our results stress, however, that both selecting the proper scale at which to assess landscape associations and empirically testing the models derived from those associations are crucial for building useful predictive models.

Landscape Ecology

Composition of cavity-nesting bird communities in montane aspen woodland fragments: The roles of landscape context and forest structure

We compared cavity-nesting bird communities in aspen ( Populus tremuloides ) woodland fragments classified on the basis of vegetation structure (tree density) and landscape context (surrounding vegetation). We found very few cavity nesters in fragments predominantly surrounded by forests. Fragments adjacent to meadows contained more species and a greater abundance of cavity nesters. Species richness and abundance were higher in sparsely than in densely treed meadow fragments. Because secondary cavity nesters are often limited by cavity availability, we augmented natural cavities with nest boxes. Although only five boxes contained bird nests, these were all in sparse aspen fragments predominantly surrounded by meadows. However, we found 25 northern flying squirrel ( Glaucomys sabrinus ) nests in boxes, none of which were in sparse meadow fragments. In addition to high-lighting the importance of landscape context in avian and mammalian habitat relationships, our results suggest that predator or competitor interactions may help structure this cavity-nester community.

Utah

Assessing map accuracy in a remotely sensed, ecoregion-scale cover map

Landscape- and ecoregion-based conservation efforts increasingly use a spatial component to organize data for analysis and interpretation. A challenge particular to remotely sensed cover maps generated from these efforts is how best to assess the accuracy of the cover maps, especially when they can exceed 1000 s/km2 in size. Here we develop and describe a methodological approach for assessing the accuracy of large-area cover maps, using as a test case the 21.9 million ha cover map developed for Utah Gap Analysis. As part of our design process, we first reviewed the effect of intracluster correlation and a simple cost function on the relative efficiency of cluster sample designs to simple random designs. Our design ultimately combined clustered and subsampled field data stratified by ecological modeling unit and accessibility (hereafter a mixed design). We next outline estimation formulas for simple map accuracy measures under our mixed design and report results for eight major cover types and the three ecoregions mapped as part of the Utah Gap Analysis. Overall accuracy of the map was 83.2% (SE=1.4). Within ecoregions, accuracy ranged from 78.9% to 85.0%. Accuracy by cover type varied, ranging from a low of 50.4% for barren to a high of 90.6% for man modified. In addition, we examined gains in efficiency of our mixed design compared with a simple random sample approach. In regard to precision, our mixed design was more precise than a simple random design, given fixed sample costs. We close with a discussion of the logistical constraints facing attempts to assess the accuracy of large-area, remotely sensed cover maps.

Remote Sensing of Environment