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Bias correction of bounded location errors in presence-only data

Location error occurs when the true location is different than the reported location. Because habitat characteristics at the true location may be different than those at the reported location, ignoring location error may lead to unreliable inference concerning species–habitat relationships. We explain how a transformation known in the spatial statistics literature as a change of support (COS) can be used to correct for location errors when the true locations are points with unknown coordinates contained within arbitrary shaped polygons. We illustrate the flexibility of the COS by modelling the resource selection of Whooping Cranes ( Grus americana ) using citizen contributed records with locations that were reported with error. We also illustrate the COS with a simulation experiment. In our analysis of Whooping Crane resource selection, we found that location error can result in up to a five-fold change in coefficient estimates. Our simulation study shows that location error can result in coefficient estimates that have the wrong sign, but a COS can efficiently correct for the bias.

Methods in Ecology and Evolution

Estimating occupancy and abundance using aerial images with imperfect detection

Species distribution and abundance are critical population characteristics for efficient management, conservation, and ecological insight. Point process models are a powerful tool for modelling distribution and abundance, and can incorporate many data types, including count data, presence-absence data, and presence-only data. Aerial photographic images are a natural tool for collecting data to fit point process models, but aerial images do not always capture all animals that are present at a site. Methods for estimating detection probability for aerial surveys usually include collecting auxiliary data to estimate the proportion of time animals are available to be detected. We developed an approach for fitting point process models using an N -mixture model framework to estimate detection probability for aerial occupancy and abundance surveys. Our method uses multiple aerial images taken of animals at the same spatial location to provide temporal replication of sample sites. The intersection of the images provide multiple counts of individuals at different times. We examined this approach using both simulated and real data of sea otters ( Enhydra lutris kenyoni ) in Glacier Bay National Park, southeastern Alaska. Using our proposed methods, we estimated detection probability of sea otters to be 0.76, the same as visual aerial surveys that have been used in the past. Further, simulations demonstrated that our approach is a promising tool for estimating occupancy, abundance, and detection probability from aerial photographic surveys. Our methods can be readily extended to data collected using unmanned aerial vehicles, as technology and regulations permit. The generality of our methods for other aerial surveys depends on how well surveys can be designed to meet the assumptions of N -mixture models.

Methods in Ecology and Evolution

Long-term video surveillance and automated analyses reveal arousal patterns in groups of hibernating bats

Understanding natural behaviours is essential to determining how animals deal with new threats (e.g. emerging diseases). However, natural behaviours of animals with cryptic lifestyles, like hibernating bats, are often poorly characterized. White-nose syndrome (WNS) is an unprecedented disease threatening multiple species of hibernating bats, and pathogen-induced changes to host behaviour may contribute to mortality. To better understand the behaviours of hibernating bats and how they might relate to WNS, we developed new ways of studying hibernation across entire seasons. We used thermal-imaging video surveillance cameras to observe little brown bats ( Myotis lucifugus ) and Indiana bats ( M. sodalis ) in two caves over multiple winters. We developed new, sharable software to test for autocorrelation and periodicity of arousal signals in recorded video. We processed 740 days (17,760 hr) of video at a rate of >1,000 hr of video imagery in less than 1 hr using a desktop computer with sufficient resolution to detect increases in arousals during midwinter in both species and clear signals of daily arousal periodicity in infected M. sodalis . Our unexpected finding of periodic synchronous group arousals in hibernating bats demonstrate the potential for video methods and suggest some bats may have innate behavioural strategies for coping with WNS. Surveillance video and accessible analysis software make it now practical to investigate long-term behaviours of hibernating bats and other hard-to-study animals.

Methods in Ecology and Evolution

Statistical design and analysis for plant cover studies with multiple sources of observation errors

Effective wildlife habitat management and conservation requires understanding the factors influencing distribution and abundance of plant species. Field studies, however, have documented observation errors in visually estimated plant cover including measurements which differ from the true value (measurement error) and not observing a species that is present within a plot (detection error). Unlike the rapid expansion of occupancy and N-mixture models for analysing wildlife surveys, development of statistical models accounting for observation error in plants has not progressed quickly. Our work informs development of a monitoring protocol for managed wetlands within the National Wildlife Refuge System. Zero-augmented beta (ZAB) regression is the most suitable method for analysing areal plant cover recorded as a continuous proportion but assumes no observation errors. We present a model extension that explicitly includes the observation process thereby accounting for both measurement and detection errors. Using simulations, we compare our approach to a ZAB regression that ignores observation errors (naïve model) and an “ad hoc” approach using a composite of multiple observations per plot within the naïve model. We explore how sample size and within-season revisit design affect the ability to detect a change in mean plant cover between 2 years using our model. Explicitly modelling the observation process within our framework produced unbiased estimates and nominal coverage of model parameters. The naïve and “ad hoc” approaches resulted in underestimation of occurrence and overestimation of mean cover. The degree of bias was primarily driven by imperfect detection and its relationship with cover within a plot. Conversely, measurement error had minimal impacts on inferences. We found >30 plots with at least three within-season revisits achieved reasonable posterior probabilities for assessing change in mean plant cover. For rapid adoption and application, code for Bayesian estimation of our single-species ZAB with errors model is included. Practitioners utilizing our R-based simulation code can explore trade-offs among different survey efforts and parameter values, as we did, but tuned to their own investigation. Less abundant plant species of high ecological interest may warrant the additional cost of gathering multiple independent observations in order to guard against erroneous conclusions.

Methods in Ecology and Evolution

Imperfect pathogen detection from non-invasive skin swabs biases disease inference

1. Conservation managers rely on accurate estimates of disease parameters, such as pathogen prevalence and infection intensity, to assess disease status of a host population. However, these disease metrics may be biased if low-level infection intensities are missed by sampling methods or laboratory diagnostic tests. These false negatives underestimate pathogen prevalence and overestimate mean infection intensity of infected individuals. 2. Our objectives were two-fold. First, we quantified false negative error rates of Batrachochytrium dendrobatidis on non-invasive skin swabs collected from an amphibian community in El Copé, Panama. We swabbed amphibians twice in sequence, and we used a recently developed hierarchical Bayesian estimator to assess disease status of the population. Second, we developed a novel hierarchical Bayesian model to simultaneously account for imperfect pathogen detection from field sampling and laboratory diagnostic testing. We evaluated the performance of the model using simulations and varying sampling design to quantify the magnitude of bias in estimates of pathogen prevalence and infection intensity. 3. We show that Bd detection probability from skin swabs was related to host infection intensity, where Bd infections < 10 zoospores have < 95% probability of being detected. If imperfect Bd detection was not considered, then Bd prevalence was underestimated by as much as 16%. In the Bd-amphibian system, this indicates a need to correct for imperfect pathogen detection caused by skin swabs in persisting host communities with low-level infections. More generally, our results have implications for study designs in other disease systems, particularly those with similar objectives, biology, and sampling decisions. 4. Uncertainty in pathogen detection is an inherent property of most sampling protocols and diagnostic tests, where the magnitude of bias depends on the study system, type of infection, and false negative error rates. Given that it may be difficult to know this information in advance, we advocate that the most cautious approach is to assume all errors are possible and to accommodate them by adjusting sampling designs. The modeling framework presented here improves the accuracy in estimating pathogen prevalence and infection intensity.

Methods in Ecology and Evolution

A new framework for analysing automated acoustic species detection data: Occupancy estimation and optimization of recordings post-processing

The development and use of automated species-detection technologies, such as acoustic recorders, for monitoring wildlife are rapidly expanding. Automated classification algorithms provide a cost- and time-effective means to process information-rich data, but often at the cost of additional detection errors. Appropriate methods are necessary to analyse such data while dealing with the different types of detection errors. We developed a hierarchical modelling framework for estimating species occupancy from automated species-detection data. We explore design and optimization of data post-processing procedures to account for detection errors and generate accurate estimates. Our proposed method accounts for both imperfect detection and false positive errors and utilizes information about both occurrence and abundance of detections to improve estimation. Using simulations, we show that our method provides much more accurate estimates than models ignoring the abundance of detections. The same findings are reached when we apply the methods to two real datasets on North American frogs surveyed with acoustic recorders. When false positives occur, estimator accuracy can be improved when a subset of detections produced by the classification algorithm is post-validated by a human observer. We use simulations to investigate the relationship between accuracy and effort spent on post-validation, and found that very accurate occupancy estimates can be obtained with as little as 1% of data being validated. Automated monitoring of wildlife provides opportunity and challenges. Our methods for analysing automated species-detection data help to meet key challenges unique to these data and will prove useful for many wildlife monitoring programs.

Methods in Ecology and Evolution

Integrating continuous stocks and flows into state-and-transition simulation models of landscape change

State-and-transition simulation models (STSMs) provide a general framework for forecasting landscape dynamics, including projections of both vegetation and land-use/land-cover (LULC) change. The STSM method divides a landscape into spatially-referenced cells and then simulates the state of each cell forward in time, as a discrete-time stochastic process using a Monte Carlo approach, in response to any number of possible transitions. A current limitation of the STSM method, however, is that all of the state variables must be discrete. Here we present a new approach for extending a STSM, in order to account for continuous state variables, called a state-and-transition simulation model with stocks and flows (STSM-SF). The STSM-SF method allows for any number of continuous stocks to be defined for every spatial cell in the STSM, along with a suite of continuous flows specifying the rates at which stock levels change over time. The change in the level of each stock is then simulated forward in time, for each spatial cell, as a discrete-time stochastic process. The method differs from the traditional systems dynamics approach to stock-flow modelling in that the stocks and flows can be spatially-explicit, and the flows can be expressed as a function of the STSM states and transitions. We demonstrate the STSM-SF method by integrating a spatially-explicit carbon (C) budget model with a STSM of LULC change for the state of Hawai'i, USA. In this example, continuous stocks are pools of terrestrial C, while the flows are the possible fluxes of C between these pools. Importantly, several of these C fluxes are triggered by corresponding LULC transitions in the STSM. Model outputs include changes in the spatial and temporal distribution of C pools and fluxes across the landscape in response to projected future changes in LULC over the next 50 years. The new STSM-SF method allows both discrete and continuous state variables to be integrated into a STSM, including interactions between them. With the addition of stocks and flows, STSMs provide a conceptually simple yet powerful approach for characterizing uncertainties in projections of a wide range of questions regarding landscape change.

Hawai'i

Two-species occupancy modeling accounting for species misidentification and nondetection

In occupancy studies, species misidentification can lead to false‐positive detections, which can cause severe estimator biases. Currently, all models that account for false‐positive errors only consider omnibus sources of false detections and are limited to single‐species occupancy. However, false detections for a given species often occur because of the misidentification with another, closely related species. To exploit this explicit source of false‐positive detection error, we develop a two‐species occupancy model that accounts for misidentifications between two species of interest. As with other false‐positive models, identifiability is greatly improved by the availability of unambiguous detections at a subset of site x occasions. Here, we consider the case where some of the field observations can be confirmed using laboratory or other independent identification methods (“confirmatory data”). We performed three simulation studies to (1) assess the model's performance under various realistic scenarios, (2) investigate the influence of the proportion of confirmatory data on estimator accuracy and (3) compare the performance of this two‐species model with that of the single‐species false‐positive model. The model shows good performance under all scenarios, even when only small proportions of detections are confirmed (e.g. 5%). It also clearly outperforms the single‐species model. We illustrate application of this model using a 4‐year dataset on two sympatric species of lungless salamanders: the US federally endangered Shenandoah salamander Plethodon shenandoah , and its presumed competitor, the red‐backed salamander Plethodon cinereus . Occupancy of red‐backed salamanders appeared very stable across the 4 years of study, whereas the Shenandoah salamander displayed substantial turnover in occupancy of forest habitats among years. Given the extent of species misidentification issues in occupancy studies, this modelling approach should help improve the reliability of estimates of species distribution, which is the goal of many studies and monitoring programmes. Further developments, to account for different forms of state uncertainty, can be readily undertaken under our general approach.

Methods in Ecology and Evolution

Evaluation of acoustic telemetry grids for determining aquatic animal movement and survival

Acoustic telemetry studies have frequently prioritized linear configurations of hydrophone receivers, such as perpendicular from shorelines or across rivers, to detect the presence of tagged aquatic animals. This approach introduces unknown bias when receivers are stationed for convenience at geographic bottlenecks (e.g., at the mouth of an embayment or between islands) as opposed to deployments following a statistical sampling design. We evaluated two-dimensional acoustic receiver arrays (grids: receivers spread uniformly across space) as an alternative approach to provide estimates of survival, movement, and habitat use. Performance of variably-spaced receiver grids (5–25 km spacing) was evaluated by simulating (1) animal tracks as correlated random walks (speed: 0.1–0.9 m/s; turning angle standard deviation: 5–30 degrees); (2) variable tag transmission intervals along each track (nominal delay: 15–300 seconds); and (3) probability of detection of each transmission based on logistic detection range curves (midpoint: 200–1500 m). From simulations, we quantified i) time between successive detections on any receiver (detection time), ii) time between successive detections on different receivers (transit time), and iii) distance between successive detections on different receivers (transit distance). In the most restrictive detection range scenario (200 m), the 95 th percentile of transit time was 3.2 days at 5 km grid spacing, 5.7 days at 7 km, and 15.2 days at 25 km; for the 1500 m detection range scenario, it was 0.1 days at 5 km, 0.5 days at 7 km, and 10.8 days at 25 km. These values represented upper bounds on the expected maximum time that an animal could go undetected. Comparison of the simulations with pilot studies on three fishes (walleye Sander vitreus , common carp Cyprinus carpio , and channel catfish Ictalurus punctatus ) from two independent large lake ecosystems (lakes Erie and Winnipeg) revealed shorter detection and transit times than what simulations predicted. By spreading effort uniformly across space, grids can improve understanding of fish migration over the commonly employed receiver line approach, but at increased time cost for maintaining grids.

Methods in Ecology and Evolution

Animal movement models for migratory individuals and groups

Animals often exhibit changes in their behaviour during migration. Telemetry data provide a way to observe geographic position of animals over time, but not necessarily changes in the dynamics of the movement process. Continuous‐time models allow for statistical predictions of the trajectory in the presence of measurement error and during periods when the telemetry device did not record the animal's position. However, continuous‐time models capable of mimicking realistic trajectories with sufficient detail are computationally challenging to fit to large datasets. Furthermore, basic continuous‐time model specifications (e.g. Brownian motion) lack realism in their ability to capture nonstationary dynamics. We present a unified class of animal movement models that are computationally efficient and provide a suite of approaches for accommodating nonstationarity in continuous trajectories due to migration and interactions among individuals. Our approach uses process convolutions to allow for flexibility in the movement process while facilitating implementation and incorporating location uncertainty. We show how to nest convolution models to incorporate interactions among migrating individuals to account for nonstationarity and provide inference about dynamic migratory networks. We demonstrate these approaches in two case studies involving migratory birds. Specifically, we used process convolution models with temporal deformation to account for heterogeneity in individual greater white‐fronted goose migrations in Europe and Iceland, and we used nested process convolutions to model dynamic migratory networks in sandhill cranes in North America. The approach we present accounts for various forms of temporal heterogeneity in animal movement and is not limited to migratory applications. Furthermore, our models rely on well‐established principles for modelling‐dependent data and leverage modern approaches for modelling dynamic networks to help explain animal movement and social interaction.

Methods in Ecology and Evolution

Using partial aggregation in spatial capture recapture

Spatial capture–recapture (SCR) models are commonly used for analysing data collected using noninvasive genetic sampling (NGS). Opportunistic NGS often leads to detections that do not occur at discrete detector locations. Therefore, spatial aggregation of individual detections into fixed detectors (e.g., centre of grid cells) is an option to increase computing speed of SCR analyses. However, it may reduce precision and accuracy of parameter estimations. Using simulations, we explored the impact that spatial aggregation of detections has on a trade‐off between computing time and parameter precision and bias, under a range of biological conditions. We used three different observation models: the commonly used Poisson and Bernoulli models, as well as a novel way to partially aggregate detections (Partially Aggregated Binary model [PAB]) to reduce the loss of information after aggregating binary detections. The PAB model divides detectors into K subdetectors and models the frequency of subdetectors with more than one detection as a binomial response with a sample size of K. Finally, we demonstrate the consequences of aggregation and the use of the PAB model using NGS data from the monitoring of wolverine ( Gulo gulo ) in Norway. Spatial aggregation of detections, while reducing computation time, does indeed incur costs in terms of reduced precision and accuracy, especially for the parameters of the detection function. SCR models estimated abundance with a low bias (<10%) even at high degree of aggregation, but only for the Poisson and PAB models. Overall, the cost of aggregation is mitigated when using the Poisson and PAB models. At the same level of aggregation, the PAB observation model out‐performs the Bernoulli model in terms of accuracy of estimates, while offering the benefits of a binary observation model (less assumptions about the underlying ecological process) over the count‐based model. We recommend that detector spacing after aggregation does not exceed 1.5 times the scale‐parameter of the detection function in order to limit bias. We recommend the use of the PAB observation model when performing spatial aggregation of binary data as it can mitigate the cost of aggregation, compared to the Bernoulli model.

Methods in Ecology and Evolution

Modelling sound attenuation in heterogeneous environments for improved bioacoustic sampling of wildlife populations

Acoustic sampling methods are becoming increasingly important in biological monitoring. Sound attenuation is one of the most important dynamics affecting the utility of acoustic data as it directly affects the probability of detection of individuals by acoustic sensor arrays and especially the localization of acoustic signals necessary in telemetry studies. Therefore, models of sound attenuation are necessary to make efficient use of acoustic data in ecological monitoring and assessment applications. Models of attenuation in widespread use are based on Euclidean distance between source and sensor, which is justified under spherical attenuation of sound waves in homogeneous environments. In this paper, I develop a model of sound attenuation based on a non‐Euclidean cost‐weighted distance metric which contains attenuation coefficients that characterize the attenuation of sound due to environmental heterogeneity in the vicinity of an acoustic sensor array. I show that parameters of the proposed attenuation model can be estimated by maximum likelihood using experimental data from an array of fixed sources, thus allowing investigators who use bioacoustic methods to devise explicit models of sound attenuation in situ and apply them to localization of sources and density estimation. In addition, drawing on analogy with spatial capture–recapture models, I argue that parameters of the non‐Euclidean model of attenuation can be estimated when source locations are unknown . Thus, the models can be applied to real field studies which require estimation of attenuation parameters or localization of signals. Models of heterogeneous sound attenuation allow more accurate descriptions of acoustic monitoring data, and therefore should produce more accurate estimates of ecological parameters of interest, including source locations, density, and movement trajectories. Moreover, the ability to test specific hypotheses about the effects of habitat and landscape structure on sound attenuation can improve the design of acoustic monitoring arrays and lead to more efficient deployment of acoustic sensing technology.

Methods in Ecology and Evolution

DNA mixtures for ecology

Mixtures of DNA from multiple contributors present a novel opportunity to count individuals to inform fish and wildlife ecology. We apply a likelihood-based framework to estimate the number of contributors to a DNA mixture for ecological applications. We then assess the performance of DNA mixture estimation through a combination of simulation analyses, laboratory testing, and a field trial to estimate fish predation rates from stomach content analysis. Simulations indicated reasonably sized genetic marker panels could estimate the number of contributors to mixtures comprised of up to 10 individuals, with potential to resolve larger mixtures with additional markers. Mixture estimates demonstrated robustness to common genotyping errors associated with fish and wildlife genetics applications. Laboratory trials demonstrated that DNA combined from multiple yellow perch ( Perca flavescens ) could be successfully genotyped with a 14-loci microsatellite panel and led to successful estimation for up to 5-contributor mixtures. Stomach content analysis with DNA mixtures indicated a 5-fold increase in estimated predation rates of yellow perch by largemouth bass ( Micropterus salmoides ) relative to conventional visual assessment of diet contents which can miss partially digested prey items. DNA mixtures have potential to expand applications of count-based ecological analyses. Technical challenges in generating genotypes from DNA mixtures may initially limit their use, however, advances in next generation genotyping platforms are anticipated to surmount these obstacles. Chiefly, we envision opportunity for DNA mixtures to advance eDNA analysis beyond presence/absence based inference to enumeration of specimens.

Methods in Ecology and Evolution

Occupancy models for citizen-science data

Large‐scale citizen‐science projects, such as atlases of species distribution, are an important source of data for macroecological research, for understanding the effects of climate change and other drivers on biodiversity, and for more applied conservation tasks, such as early‐warning systems for biodiversity loss. However, citizen‐science data are challenging to analyse because the observation process has to be taken into account. Typically, the observation process leads to heterogeneous and non‐random sampling, false absences, false detections, and spatial correlations in the data. Increasingly, occupancy models are being used to analyse atlas data. We advocate a dual approach to strengthen inference from citizen science data for the questions the programme is intended to address: (a) the survey design should be chosen with a particular set of questions and associated analysis strategy in mind and (b) the statistical methods should be tailored not only to those questions but also to the specific characteristics of the data. We review the consequences of particular survey design choices that typically need to be made in atlas‐style citizen‐science projects. These include spatial resolution of the sampling units, allocation of effort in space, and collection of information about the observation process. On the analysis side, we review extensions of the basic occupancy models that are frequently necessary with atlas data, including methods for dealing with heterogeneity, non‐independent detections, false detections, and violation of the closure assumption. New technologies, such as cell‐phone apps and fixed remote detection devices, are revolutionizing citizen‐science projects. There is an opportunity to maximize the usefulness of the resulting datasets if the protocols are rooted in robust statistical designs and data analysis issues are being considered. Our review provides guidelines for designing new projects and an overview of the current methods that can be used to analyse data from such projects.

Methods in Ecology and Evolution

The area under the precision‐recall curve as a performance metric for rare binary events

Species distribution models are used to study biogeographic patterns and guide decision‐making. The variable quality of these models makes it critical to assess whether a model's outputs are suitable for the intended use, but commonly used evaluation approaches are inappropriate for many ecological contexts. In particular, unrealistically high performance assessments have been associated with models for rare species and predictions over large geographic extents. We evaluated the area under the precision‐recall curve (AUC‐PR) as a performance metric for rare binary events, focusing on the assessment of species distribution models. Precision is the probability that a species is present given a predicted presence, while recall (more commonly called sensitivity) is the probability the model predicts presence in locations where the species has been observed. We simulated species at three levels of prevalence, compared AUC‐PR and the area under the receiver operating characteristic curve (AUC‐ROC) when the geographic extent of predictions was increased and assessed how well each metric reflected a model's utility to guide surveys for new populations. AUC‐PR was robust to species rarity and, unlike AUC‐ROC, not affected by an increasing geographic extent. The major advantages of AUC‐PR arise because it does not incorporate correctly predicted absences and is therefore less prone to exaggerate model performance for unbalanced datasets. AUC‐PR and precision were useful indicators of a model's utility for guiding surveys. We show that AUC‐PR has important advantages for evaluating models of rare species, and its benefits in the context of unbalanced binary responses will make it applicable for other ecological studies. By not considering the true negative quadrant of the confusion matrix, AUC‐PR ameliorates issues that arise when the geographic extent is increased beyond the species’ range or when a large number of background points are used when absence information is unavailable. However, no single metric captures all aspects of performance nor provides an absolute index that can be compared across datasets. Our results indicate AUC‐PR and precision can provide useful and intuitive metrics for evaluating a model's utility for guiding sampling, and can complement other metrics to help delineate a model's appropriate use.

Methods in Ecology and Evolution

A comparative analysis of common methods to identify waterbird hotspots

1. Hotspot analysis is a commonly used method in ecology and conservation to identify areas of high biodiversity or conservation concern. However, delineating and mapping hotspots is subjective and various approaches can lead to different conclusions with regard to the classification of particular areas as hotspots, complicating long-term conservation planning and implementation efforts. 2. We present a comparative analysis of recent approaches for identifying waterbird hotspots, with the goal of developing insights about the appropriate use of these methods. We selected four commonly used measures to identify persistent areas of high use: kernel density estimation, Getis-Ord Gi*, hotspot persistence, and hotspots conditional on presence, which represent the range of quantitative hotspot estimation approaches used in waterbird analyses. We applied each of the methods to aerial survey waterbird count data collected in the Great Lakes from 2012-2014 using a 5 km2 grid. For each approach, we identified areas of high use for seven species/species groups and then compared the results across all methods. 3. Our results indicate that formal hotspot analysis frameworks do not always lead to the same conclusions. The kernel density and Getis-Ord Gi* methods yielded the most similar results across all species analyzed. We found that these two models can differ substantially from the hotspot persistence and hotspots conditional on presence estimation approaches, which were not consistently similar to one another. The hotspot persistence approach differed most significantly from the other methods but is the only method to explicitly account for temporal variation. 4. We recommend considering the ecological question and scale of any conservation or management activities prior to designing survey methodologies. Deciding the appropriate definition and scale for analysis is critical for interpretation of hotspot analysis results. Combining methods using an integrative approach, either within a single analysis or post-hoc, could lead to greater consistency in the identification of waterbird hotspots.

Lake Erie, Lake Huron, Lake Michigan

Cohesive framework for modeling plant cover class data

The study of plant distribution and abundance is a fundamental pursuit in ecology and conservation biology. Measuring plant abundance by visually assessing percent cover and recording a cover class is a common field method that yields ordinal data. Statistical models for ordinal data exist but entail cumbersome interpretations and sometimes restrictive assumptions. We propose a Bayesian hierarchical framework for analysing cover class data that allows for linking ordinal observations to a latent beta distribution and accounts for zero inflation. Harnessing a latent beta distribution supports interpreting changes in abundance in terms of mean percent cover rather than odds ratios of cumulative cover classes as for cumulative link models. The zero augmentation allows for simultaneous inferences on both occurrence (distribution) and abundance. We show how our model can account for true and false zeros, misclassification of cover classes, multiple species and hierarchical sampling designs, using empirical examples and simulations. Simulated observation errors, when ignored, led to models overestimating abundance and underestimating occurrence. Based on simulations, we found no substantial difference between mean percent cover estimates when analyzing ordinal cover classes versus continuous percent cover as the response. Our empirical datasets displayed high probability of detection (>0.85 on average for all species), likely due to the sampling design used and training of observers. Probability of occurrence was slightly underestimated for bare ground, Artemisia tridentata , Elycap medusae , and Poa secunda using a model that ignored imperfect detection. Estimated mean percent cover was not substantially impacted by ignoring measurement error for five plant species and bare ground. Our modelling framework for cover class data allows for an explicit separation of distribution from abundance and, importantly, allows for interpreting species–environment relationships in terms of variation in mean percent cover as compared to cumulative odds ratios. The beta distribution inherently accommodates heteroscedasticity and skewness, statistical properties that are a consequence of spatially aggregated patterns common to plant survey data. Recording cover classes provides a reliable, efficient way to measure plants and our simulations suggest little loss of information compared to assuming continuous percent cover. We provide JAGS and Stan model code for implementation.

Methods in Ecology and Evolution

Sampling and analysis frameworks for inference in ecology

1. Reliable statistical inference is central to ecological research, much of which seeks to estimate population attributes and their interactions. The issue of sampling design and its relationship to inference has become increasingly important due to rapid proliferation of modeling methodology (line transect modeling, capture-recapture, estimation of occurrence, model selection procedures, hierarchical modeling) and new sampling approaches (adaptive sampling, other specialized designs). It is important for ecologists using these advanced methods to be aware of how the linkages between sample selection and data analysis can potentially affect inference. 2. We examine design-based and model-based inference frameworks for ecological data collected randomly, purposively, or opportunistically. We elucidate differences in the probability structures for data arising from these frameworks, clarify the assumptions that underlie them, and demonstrate their differences. 3. Design-based inference builds on a probability structure inherited from randomized data collection, whereas model-based inference relies on an assumed stochastic model of the data. By itself, a design-based approach is of limited value for inferences about causal hypotheses. In contrast, model-based inference is dependent on a conditionality principle that can seldom be shown to be met for an ecological system. We describe the conditions under which one can safely ignore sampling design in model-based analysis, along with inferential implications if these conditions are not met. The special case of opportunistic sampling is discussed. 4. We present a combined framework that takes advantage of both approaches to inference, and provides a robust methodology that can deal with the modeling of sampling problems such as nondetection and misclassification, as well as the exploration of causal hypotheses. The combined framework can be useful for identifying optimal sampling strategies. 5. Each approach to inference has its strengths and weaknesses, and practitioners should be aware of these in order to tailor designs and analyses to specific questions. We use the approaches and their underlying rationales to provide guidelines for choosing designs and estimators for reliable inference.

Methods in Ecology and Evolution