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Propagating observation errors to enable scalable and rigorous enumeration of plant population abundance with aerial imagery

Estimating and monitoring plant population size is fundamental for ecological research, as well as conservation and restoration programs. High-resolution imagery has potential to facilitate such estimation and monitoring. However, remotely sensed estimates typically have higher uncertainty than field measurements, risking biased inference on population status. We present a model that accounts for false negative (missed plants) and false positive (misclassified or double-counted plants) error in counts from high-resolution imagery via integration with ground data. We apply it to estimate the abundance of a foundational shrub species in post-wildfire landscapes in the western United States. In these landscapes, plant recruitment is crucial for ecological recovery but locally patchy, motivating the use of spatially extensive measurements from unoccupied aerial systems (UAS). Integrating >16 ha of UAS imagery with >700 georeferenced field plots, we fit our model to generate insights into the prevalence and drivers of observation errors associated with classification algorithms used to distinguish individual plants, relationships between abundance and landscape context, and to generate spatially explicit maps of shrub abundance. Raw counts of plant abundance in high-resolution imagery resulted in substantial false negative and false positive observation errors. The probability of detecting ( p ) adult plants (≥0.25 m tall) varied between sites within 0.52 < p̂adult < 0.82, whereas the detection of smaller plants (<0.25 m) was lower, 0.03 < p̂small < 0.3. On average, we estimate that 19% of all detected plants were false positive errors, which varied spatially in relation to topographic predictors. Abundance declined toward the interior of previous wildfires and was positively associated with terrain roughness. Our study demonstrates that integrated models accounting for imperfect detection improve estimates of plant population abundance derived from inherently imperfect UAS imagery. We believe such models will further improve inference on plant population dynamics—relevant to restoration, wildlife habitat and related objectives—and echo previous calls for remote sensing applications to better differentiate between ecological and observational processes.

Methods in Ecology and Evolution↗

Detecting stochasticity in population time series using a non-parametric test of intrinsic predictability

1. Many ecological systems dominated by stochastic dynamics can produce complex time series that inherently limit forecast accuracy. The ‘intrinsic predictability’ of these systems can be approximated by a time series complexity metric called weighted permutation entropy (WPE). While WPE is a useful metric to gauge forecast performance prior to model building, it is sensitive to noise and may be biased depending on the length of the time series. Here, we introduce a simple randomized permutation test (rWPE) to assess whether a time series is intrinsically more predictable than white noise. 2. We apply rWPE to both simulated and empirical data to assess its performance and usefulness. To do this, we simulate population dynamics under various scenarios, including a linear trend, chaotic, periodic and equilibrium dynamics. We further test this approach with observed abundance time series for 932 species across four orders of animals from the Global Population Dynamics Database. Finally, using Adélie ( Pygoscelis adeliae ) and emperor penguin ( Aptenodytes forsteri ) time series as case studies, we demonstrate the application of rWPE to multiple populations for a single species. 3. We show that rWPE can determine whether a system is significantly more predictable than white noise, even with time series as short as 10 years that show an apparent trend under biologically realistic stochasticity levels. Additionally, rWPE has statistical power close to 100% when time series are at least 30 time steps long and show chaotic or periodic dynamics. Power decreases to ~10% under equilibrium dynamics, irrespective of time series length. Among four classes of animal taxa, mammals have the highest relative frequency (28%) of time series that are both longer than 30 time steps and indistinguishable from white noise in terms of complexity, followed by insects (16%), birds (16%) and bony fishes (11%). 4. rWPE is a straightforward and useful method widely applicable to any time series, including short ones. By informing forecasters of the inherent limitations to a system's predictability, it can guide a modeller's expectations for forecast performance.

Methods in Ecology and Evolution↗

Thinking beyond the closure assumption: Designing surveys for estimating biological truth with occupancy models

Occupancy models estimate distributions of imperfectly detected species, but violations of the closure assumption can bias results. However, researchers working with mobile animals may find it impossible to eliminate such violations. Here, we tested the hypothesis that occupancy models fit to realistic sampling data can generate unbiased occupancy estimates for an itinerant Wood Thrush ( Hylocichla mustelina ) population. In 2013 and 2014, we tracked movements of 41 breeding Wood Thrush males. We modelled territory shift probabilities using logistic exposure models and within-territory movements using continuous-time stochastic process models. We then constructed an individual-based model, simulated (1000 iterations) spatiotemporal locations for individuals and simulated sampling these populations using 162 different point count protocols with variable spatial (sampling radius and point placement method), and temporal (survey length, between-survey intervals and number of surveys) characteristics. We compared occupancy estimates with true values of instantaneous, daily and seasonal occupancy from the simulations. We parameterized continuous time stochastic process models based on movements within 34 unique territories and estimated a daily territory shift probability of 0.0099 (95% CI: 0.0060, 0.0152). Simulated data indicated that estimates of occupancy ranged from 0.18 (0.06, 1.00) to 0.80 (0.71, 0.89) depending on protocol characteristics. Occupancy estimates increased with increasing survey radius, survey length and between-survey interval. Protocols using shorter surveys and between-survey intervals were good estimators for instantaneous occupancy (low bias and mean-squared error) but poor estimators for daily and seasonal occupancy; longer surveys and intervals generated unbiased estimators of daily occupancy but underestimated seasonal occupancy. Logistic regression models that ignored imperfect detection outperformed occupancy models for estimating instantaneous occupancy but not daily or seasonal occupancy. For mobile animals, occupancy of sampling sites changes in space and time. Consequently, the spatial and temporal aspects of a sampling protocol have strong, but predictable, effects on occupancy model parameter estimates. Our results demonstrate that how these factors interact is critical for designing surveys that produce occupancy estimates representative of the biological process of interest to a researcher.

Methods in Ecology and Evolution↗

A transferable approach for quantifying benthic fish sizes and densities in annotated underwater images

1. Benthic fishes are a common target of scientific monitoring but are difficult to quantify because of their close association to bottom habitats that are hard to access. Advances in image-acquisition technologies, machine vision, and deep learning have made capturing and quantifying fishes with cameras increasingly feasible. We present a method and open-source software called ‘FishScale’ to estimate benthic fish lengths, numeric abundance, and biomass density in underwater environments assessed with down-looking monocular images. 2. ‘FishScale’ estimates fish abundances and size frequencies from near-nadir monocular images where fish have already been semantically segmented. The software accounts for lens distortion, underwater magnification effects, and fish body curvature to automatically estimate fish lengths and the areas of images where they were captured. Numeric and biomass density are estimated through a deterministic machine vision algorithm that requires a user-provided length-weight relationship for species of interest and calibration images. 3. Results from validation studies show that lengths and weights can be estimated with high accuracy and precision for round goby ( Neogobius melanostomus ) captured in distorted action camera images, and from large-bodied lake trout ( Salvelinus namaycush ) imaged with a machine vision camera. The real-world utility of the approach is demonstrated in a case study estimating round goby abundances and size frequencies along a 10.7-km transect surveyed with an autonomous underwater vehicle in Lake Michigan, USA. 4. Our validation studies demonstrate that the approach estimates benthic and benthopelagic fish lengths and weights with little bias and good accuracy and precision for species with much different body shapes and sizes. The method is applicable to data collected using a variety of nadir imaging approaches with widespread applications to fisheries monitoring and quantification of any species or object for which nadir images and working distances between the camera and feature of interest are available.

Illinois, Indiana, Michigan, Wisconsin↗

Challenges and opportunities for data integration to improve estimation of migratory connectivity

Understanding migratory connectivity, or the linkage of populations between seasons, is critical for effective conservation and management of migratory wildlife. A growing number of tools are available for understanding where migratory individuals and populations occur throughout the annual cycle. Integration of the diverse measures of migratory movements can help elucidate migratory connectivity patterns with methodology that accounts for differences in sampling design, directionality, effort, precision and bias inherent to each data type. The R package MigConnectivity was developed to estimate population-specific connectivity and the range-wide strength of those connections. New functions allow users to integrate intrinsic markers, tracking and long-distance reencounter data, collected from the same or different individuals, to estimate population-specific transition probabilities (estTransition) and the range-wide strength of those transition probabilities (estStrength). We used simulation and real-world case studies to explore the challenges and limitations of data integration based on data from three migratory bird species, Painted Bunting ( Passerina ciris ), Yellow Warbler ( Setophaga petechia ) and Bald Eagle ( Haliaeetus leucocephalus ), two of which had bidirectional data. We found data integration is useful for quantifying migratory connectivity, as single data sources are less likely to be available across the species range. Furthermore, accurate strength estimates can be obtained from either breeding-to-nonbreeding or nonbreeding-to-breeding data. For bidirectional data, integration can lead to more accurate estimates when data are available from all regions in at least one season. The ability to conduct combined analyses that account for the unique limitations and biases of each data type is a promising possibility for overcoming the challenge of range-wide coverage that has been hard to achieve using single data types. The best-case scenario for data integration is to have data from all regions, especially if the question is range-wide or data are bidirectional. Multiple data types on animal movements are becoming increasingly available and integration of these growing datasets will lead to a better understanding of the full annual cycle of migratory animals.

Methods in Ecology and Evolution↗

Sampling mass mortality events to enable diagnoses: A protocol using freshwater mussels

Many taxa around the globe are threatened by often unexplained mass mortality events (MMEs), which can decimate populations and compromise key ecosystem functions. One example of a highly threatened taxon facing frequent MMEs is freshwater mussels (Unionida). There has been a recent increase in interest in understanding the causes of freshwater mussel MMEs, but standardised methodologies for how best to respond to them to facilitate diagnoses are unavailable. When an MME is observed, swift and appropriate sample collection is imperative owing to the transient nature of these phenomena. Here we provide structured guidance that will facilitate rapid and appropriate sampling of MMEs, using freshwater mussels as an example. We set out standardised procedures for sample collection, preparation and preservation. The procedures we outline will improve our capacity for diagnostic investigations of MMEs and other mortality events, not only in freshwater mussels but also across many other taxa. This, in turn, can inform appropriate management responses.

Methods in Ecology and Evolution↗

Spatial close-kin mark-recapture models applied to terrestrial species with continuous natal dispersal

Close-kin mark–recapture (CKMR) methods use information on genetic relatedness among individuals to estimate demographic parameters. An individual's genotype can be considered a ‘recapture’ of each of its parent's genotype, and the frequency of kin-pair matches detected in a population sample can directly inform estimates of abundance. CKMR inference procedures require analysts to define kinship probabilities in functional forms, which inevitably involve simplifying assumptions. Among others, population structure can have a strong influence on how kinship probabilities are formulated. Many terrestrial species are philopatric or face barriers to dispersal, and not accounting for dispersal limitation in kinship probabilities, can create substantial bias if sampling is also spatially structured (e.g. via harvest). We present a spatially explicit formulation of CKMR that corrects for incomplete mixing by incorporating natal dispersal distances and spatial distribution of individuals into the kinship probabilities. We used individual-based simulations to evaluate the accuracy of abundance estimates obtained with one spatially naïve and two spatially explicit CKMR models across six scenarios with distinct spatial patterns of relative abundance and sampling probability. Estimates of abundance obtained with a CKMR model naïve to spatial structure were negatively biased when sampling was spatially biased. Incorporating patterns of natal dispersal in the kinship probabilities helped address this bias, but estimates were not always accurate depending on the model used and the scenario considered. Incorporating natal dispersal into spatially structured CKMR models can address the bias created by population structure and heterogeneous sampling but will often require additional assumptions and auxiliary data (e.g. relative abundance indices). The models shown here were designed for terrestrial species with continuous patterns of natal dispersal and high year-to-year site fidelity but could be extended to other species.

Methods in Ecology and Evolution↗

Simulated soundscapes and transfer learning boost the performance of acoustic classifiers under data scarcity

1. The biodiversity crisis necessitates spatially extensive methods to monitor multiple taxonomic groups for evidence of change in response to evolving environmental conditions. Programs that combine passive acoustic monitoring and machine learning are increasingly used to meet this need. These methods require large, annotated datasets, which are time-consuming and expensive to produce, creating potential barriers to adoption in data- and funding-poor regions. Recently released pre-trained avian acoustic classification models provide opportunities to reduce the need for manual labelling and accelerate the development of new acoustic classification algorithms through transfer learning. Transfer learning is a strategy for developing algorithms under data scarcity that uses pre-trained models from related tasks to adapt to new tasks. 2. Our primary objective was to develop a transfer learning strategy using the feature embeddings of a pre-trained avian classification model to train custom acoustic classification models in data-scarce contexts. We used three annotated avian acoustic datasets to test whether transfer learning and soundscape simulation-based data augmentation could substantially reduce the annotated training data necessary to develop performant custom acoustic classifiers. We also conducted a sensitivity analysis for hyperparameter choice and model architecture. We then assessed the generalizability of our strategy to increasingly novel non-avian classification tasks. 3. With as few as two training examples per class, our soundscape simulation data augmentation approach consistently yielded new classifiers with improved performance relative to the pre-trained classification model and transfer learning classifiers trained with other augmentation approaches. Performance increases were evident for three avian test datasets, including single-class and multi-label contexts. We observed that the relative performance among our data augmentation approaches varied for the avian datasets and nearly converged for one dataset when we included more training examples. 4. We demonstrate an efficient approach to developing new acoustic classifiers leveraging open-source sound repositories and pre-trained networks to reduce manual labelling. With very few examples, our soundscape simulation approach to data augmentation yielded classifiers with performance equivalent to those trained with many more examples, showing it is possible to reduce manual label-ling while still achieving high-performance classifiers and, in turn, expanding the potential for passive acoustic monitoring to address rising biodiversity monitoring needs.

Methods in Ecology and Evolution↗

Spatially realistic dynamic N-mixture models

Dispersal is a complex ecological process that can be driven by the abiotic environment, density-dependent processes and species mobility. Traditionally, dispersal has been studied with capture-based data that are often limited in space and time. More recently, spatial dynamic N-mixture models (i.e. N-mixture models accounting for dispersal among local populations and changes in abundance over time) allow the study of dispersal with count data. However, current spatial dynamic N-mixture models focus mainly on distance-based dispersal, limiting our ability to evaluate realistic patterns and drivers of dispersal at relevant spatial and temporal scales. We developed a spatially realistic dynamic N-mixture model that allows dispersal to be driven by environmental conditions and population density. Using simulations, we evaluated the inferential performance of this model when large proportions of sites are unsurveyed and the survey period is short, situations that are typical of real-world studies. We then used two case studies to illustrate the applications of this model, one representing a species with limited mobility in fragmented habitat (Chiricahua Leopard Frogs, Lithobates chiricahuensis , in isolated ponds) and the other representing a species with great mobility in continuous heterogeneous habitat (Common Nighthawks, Chordeiles minor , in grasslands). Our simulation study showed that the model provides valid inference even when the number of surveyed sites is small (𝑛 =10) or when the survey period is short for dynamic models (5 years), indicating that the model could have great practical value in real-world studies. The case study of Chiricahua Leopard Frogs showed that the model is capable of revealing pond-type-specific dispersal and predicting dispersal among ponds. The case study of Common Nighthawks showed the ability of the model to reveal the effect of primary productivity on dispersal and to map dispersal across the study area. Our effort demonstrates the potential of using count data to gain insights about complex yet realistic dispersal processes. The model that we introduce is flexible and adaptable for a broad range of situations in population and conservation ecology

Arizona, Kansas↗

A systematic review of avian influenza virus transmission models: Gaps and opportunities

Highly pathogenic avian influenza virus (HPAIV) increasingly threatens wild birds, poultry, non-human mammals and humans, yet gaps and opportunities in avian influenza virus (AIV) transmission modelling have not been comprehensively synthesized. We reviewed 182 articles published through 2025, categorized dimensions, including modelling approaches, data, host types, biological levels, spatial scales, temporal scales and model objectives, examined associations among these dimensions, and synthesized author-stated limitations and recommendations. Statistical and mechanistic models dominated. Model data centred on population dynamics and demography, disease surveillance, host epidemiology and competence, meteorology and land use and land cover. Most studies focused on poultry or wild birds at regional spatial scales, seasonal/annual and multi-year temporal scales, population or multi-population levels, and were retrospective. Machine learning and decision- making models were less common. Immunity, behaviour and life history, community composition, morphology, viral genetics and viral persistence received less attention. Non-human mammals, individual and community biological levels, local and broader spatial scales, daily temporal scales and prospective modelling were also less frequently modelled. The authors frequently reported limitations related to data availability, surveillance sampling and reporting bias, and model simplification, while commonly recommending improved data quality and greater model complexity and realism.

Methods in Ecology and Evolution↗

Unbiased survival estimates and evidence for skipped breeding opportunities in females

1. &ensp;Estimates of demographic parameters for females, in many organisms, are sparse. This is particularly worrisome as more and more species are faced with high extinction probabilities and conservation increasingly depends on actions dictated by complex predictive models that require accurate estimates of demographic parameters for each sex and species. 2. &ensp;This study assesses demographic parameters, specifically temporary emigration and survival, for females, a class that has been difficult to investigate historically because of lack of data. Amphibians provide a particularly good example because there is global concern about amphibian decline; yet most demographic parameter estimates are based on data from males, which we show can lead to erroneous conclusions. 3. &ensp;We use 10 years of capture&ndash;recapture data from boreal toads ( Bufo boreas ) and the multi-state open robust design model to provide evidence for the occurrence of skipped breeding opportunities (i.e. temporary emigration) in females. This is the first time that the open robust design model has been applied to an analysis of an amphibian population that we are aware of. 4. &ensp;We determined that the transition from breeder to non-breeder is obligate and the probability of a non-breeder remaining a non-breeder is 64%; thus, temporary emigration is first-order Markovian in nature, where breeding probability is dependent on the previous year&rsquo;s activity, i.e. if a female did not breed in year one, there is a 36% chance that she will breed in year two. With temporary emigration accounted for, we estimated between-year female survival at 87%. 5. &ensp;Establishing the occurrence of temporary emigration not only reduces bias in estimates of survival probabilities but also provides information about expected breeding attempts by females, a critical element in understanding the ecology of an organism and the impacts of outside stressors and conservation actions.

Methods in Ecology and Evolution↗

Assessing conservation relevance of organism-environment relations using predicted changes in response variables

1. Organism–environment models are used widely in conservation. The degree to which they are useful for informing conservation decisions – the conservation relevance of these relations – is important because lack of relevance may lead to misapplication of scarce conservation resources or failure to resolve important conservation dilemmas. Even when models perform well based on model fit and predictive ability, conservation relevance of associations may not be clear without also knowing the magnitude and variability of predicted changes in response variables. 2. We introduce a method for evaluating the conservation relevance of organism–environment relations that employs confidence intervals for predicted changes in response variables. The confidence intervals are compared to a preselected magnitude of change that marks a threshold (trigger) for conservation action. To demonstrate the approach, we used a case study from the Chihuahuan Desert involving relations between avian richness and broad-scale patterns of shrubland. We considered relations for three winters and two spatial extents (1- and 2-km-radius areas) and compared predicted changes in richness to three thresholds (10%, 20% and 30% change). For each threshold, we examined 48 relations. 3. The method identified seven, four and zero conservation-relevant changes in mean richness for the 10%, 20% and 30% thresholds respectively. These changes were associated with major (20%) changes in shrubland cover, mean patch size, the coefficient of variation for patch size, or edge density but not with major changes in shrubland patch density. The relative rarity of conservation-relevant changes indicated that, overall, the relations had little practical value for informing conservation decisions about avian richness. 4. The approach we illustrate is appropriate for various response and predictor variables measured at any temporal or spatial scale. The method is broadly applicable across ecological environments, conservation objectives, types of statistical predictive models and levels of biological organization. By focusing on magnitudes of change that have practical significance, and by using the span of confidence intervals to incorporate uncertainty of predicted changes, the method can be used to help improve the effectiveness of conservation efforts.

Methods in Ecology and Evolution↗

Accounting for non-independent detection when estimating abundance of organisms with a Bayesian approach

Summary 1. Binomial mixture models use repeated count data to estimate abundance. They are becoming increasingly popular because they provide a simple and cost‐effective way to account for imperfect detection. However, these models assume that individuals are detected independently of each other. This assumption may often be violated in the field. For instance, manatees ( Trichechus manatus latirostris ) may surface in turbid water (i.e. become available for detection during aerial surveys) in a correlated manner (i.e. in groups). However, correlated behaviour, affecting the non‐independence of individual detections, may also be relevant in other systems (e.g. correlated patterns of singing in birds and amphibians). 2. We extend binomial mixture models to account for correlated behaviour and therefore to account for non‐independent detection of individuals. We simulated correlated behaviour using beta‐binomial random variables. Our approach can be used to simultaneously estimate abundance, detection probability and a correlation parameter. 3. Fitting binomial mixture models to data that followed a beta‐binomial distribution resulted in an overestimation of abundance even for moderate levels of correlation. In contrast, the beta‐binomial mixture model performed considerably better in our simulation scenarios. We also present a goodness‐of‐fit procedure to evaluate the fit of beta‐binomial mixture models. 4. We illustrate our approach by fitting both binomial and beta‐binomial mixture models to aerial survey data of manatees in Florida. We found that the binomial mixture model did not fit the data, whereas there was no evidence of lack of fit for the beta‐binomial mixture model. This example helps illustrate the importance of using simulations and assessing goodness‐of‐fit when analysing ecological data with N‐mixture models. Indeed, both the simulations and the goodness‐of‐fit procedure highlighted the limitations of the standard binomial mixture model for aerial manatee surveys. 5. Overestimation of abundance by binomial mixture models owing to non‐independent detections is problematic for ecological studies, but also for conservation. For example, in the case of endangered species, it could lead to inappropriate management decisions, such as downlisting. These issues will be increasingly relevant as more ecologists apply flexible N‐mixture models to ecological data.

Methods in Ecology and Evolution↗

Spatial capture-recapture models for search-encounter data

1. Spatial capture&ndash;recapture models make use of auxiliary data on capture location to provide density estimates for animal populations. Previously, models have been developed primarily for fixed trap arrays which define the observable locations of individuals by a set of discrete points. 2. Here, we develop a class of models for 'search-encounter' data, i.e. for detections of recognizable individuals in continuous space, not restricted to trap locations. In our hierarchical model, detection probability is related to the average distance between individual location and the survey path. The locations are allowed to change over time owing to movements of individuals, and individual locations are related formally by a model describing individual activity or home range centre which is itself regarded as a latent variable in the model. We provide a Bayesian analysis of the model in WinBUGS, and develop a custom MCMC algorithm in the R language. 3. The model is applied to simulated data and to territory mapping data for the Willow Tit from the Swiss Breeding Bird Survey MHB. While the observed density was 15 territories per nominal 1 km 2 plot of unknown effective sample area, the model produced a density estimate of 21∙12 territories per square km (95% posterior interval: 17&ndash;26). 4. Spatial capture&ndash;recapture models are relevant to virtually all animal population studies that seek to estimate population size or density, yet existing models have been proposed mainly for conventional sampling using arrays of traps. Our model for search-encounter data, where the spatial pattern of searching can be arbitrary and may change over occasions, greatly expands the scope and utility of spatial capture&ndash;recapture models.

Methods in Ecology and Evolution↗

Applying additive modeling and gradient boosting to assess the effects of watershed and reach characteristics on riverine assemblages

Issues with ecological data (e.g. non-normality of errors, nonlinear relationships and autocorrelation of variables) and modelling (e.g. overfitting, variable selection and prediction) complicate regression analyses in ecology. Flexible models, such as generalized additive models (GAMs), can address data issues, and machine learning techniques (e.g. gradient boosting) can help resolve modelling issues. Gradient boosted GAMs do both. Here, we illustrate the advantages of this technique using data on benthic macroinvertebrates and fish from 1573 small streams in Maryland, USA.

Methods in Ecology and Evolution↗

On thinning of chains in MCMC

1. Markov chain Monte Carlo (MCMC) is a simulation technique that has revolutionised the analysis of ecological data, allowing the fitting of complex models in a Bayesian framework. Since 2001, there have been nearly 200 papers using MCMC in publications of the Ecological Society of America and the British Ecological Society, including more than 75 in the journal Ecology and 35 in the Journal of Applied Ecology . 2. We have noted that many authors routinely 'thin' their simulations, discarding all but every k th sampled value; of the studies we surveyed with details on MCMC implementation, 40% reported thinning. 3. Thinning is often unnecessary and always inefficient, reducing the precision with which features of the Markov chain are summarised. The inefficiency of thinning MCMC output has been known since the early 1990's, long before MCMC appeared in ecological publications. 4. We discuss the background and prevalence of thinning, illustrate its consequences, discuss circumstances when it might be regarded as a reasonable option and recommend against routine thinning of chains unless necessitated by computer memory limitations.

Methods in Ecology and Evolution↗

Likelihood analysis of species occurrence probability from presence-only data for modelling species distributions

1. Understanding the factors affecting species occurrence is a pre-eminent focus of applied ecological research. However, direct information about species occurrence is lacking for many species. Instead, researchers sometimes have to rely on so-called presence-only data (i.e. when no direct information about absences is available), which often results from opportunistic, unstructured sampling. MAXENT is a widely used software program designed to model and map species distribution using presence-only data. 2. We provide a critical review of MAXENT as applied to species distribution modelling and discuss how it can lead to inferential errors. A chief concern is that MAXENT produces a number of poorly defined indices that are not directly related to the actual parameter of interest &ndash; the probability of occurrence ( &psi; ). This focus on an index was motivated by the belief that it is not possible to estimate &psi; from presence-only data; however, we demonstrate that &psi; is identifiable using conventional likelihood methods under the assumptions of random sampling and constant probability of species detection. 3. The model is implemented in a convenient r package which we use to apply the model to simulated data and data from the North American Breeding Bird Survey. We demonstrate that MAXENT produces extreme under-predictions when compared to estimates produced by logistic regression which uses the full (presence/absence) data set. We note that MAXENT predictions are extremely sensitive to specification of the background prevalence, which is not objectively estimated using the MAXENT method. 4. As with MAXENT, formal model-based inference requires a random sample of presence locations. Many presence-only data sets, such as those based on museum records and herbarium collections, may not satisfy this assumption. However, when sampling is random, we believe that inference should be based on formal methods that facilitate inference about interpretable ecological quantities instead of vaguely defined indices.

Methods in Ecology and Evolution↗

A general theory of multimetric indices and their properties

1. Stewardship of biological and ecological resources requires the ability to make integrative assessments of ecological integrity. One of the emerging methods for making such integrative assessments is multimetric indices (MMIs). These indices synthesize data, often from multiple levels of biological organization, with the goal of deriving a single index that reflects the overall effects of human disturbance. Despite the widespread use of MMIs, there is uncertainty about why this approach can be effective. An understanding of MMIs requires a quantitative theory that illustrates how the properties of candidate metrics relates to MMIs generated from those metrics. 2. We present the initial basis for such a theory by deriving the general mathematical characteristics of MMIs assembled from metrics. We then use the theory to derive quantitative answers to the following questions: Is there an optimal number of metrics to comprise an index? How does covariance among metrics affect the performance of the index derived from those metrics? And what are the criteria to decide whether a given metric will improve the performance of an index? 3. We find that the optimal number of metrics to be included in an index depends on the theoretical distribution of signal of the disturbance gradient contained in each metric. For example, if the rank-ordered parameters of a metric-disturbance regression can be described by a monotonically decreasing function, then an optimum number of metrics exists and can often be derived analytically. We derive the conditions by which adding a given metric can be expected to improve an index. 4. We find that the criterion defining such conditions depends nonlinearly of the signal of the disturbance gradient, the noise (error) of the metric and the correlation of the metric errors. Importantly, we find that correlation among metric errors increases the signal required for the metric to improve the index. 5. The theoretical framework presented in this study provides the basis for understanding the properties of MMIs. It can also be useful throughout the index construction process. Specifically, it can be used to aid understanding of the benefits and limitations of combining metrics into indices; it can inform selection/collection of candidate metrics; and it can be used directly as a decision aid in effective index construction.

Methods in Ecology and Evolution↗