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Richard B. Chandler

Publications and source records attributed to Richard B. Chandler.

At least 19 recordsLinked to original sources

The unmarked R package: Twelve years of advances in occurrence and abundance modelling in ecology

Species distribution models (SDMs) are widely applied to understand the processes governing spatial and temporal variation in species abundance and distribution but often do not account for measurement errors such as false negatives and false positives. We describe unmarked , a package for the freely available and open-source R software that provides a complete workflow for modelling species distribution and abundance while explicitly accounting for measurement errors. Here we focus on recent advances in unmarked functionality to support multi-species, multi-state, and multi-season data, as well as support for fitting models with random effects. For illustration, we present an analysis of Acadian Flycatcher Empidonax virescens abundance on Roanoke River National Wildlife Refuge, North Carolina, USA, over 18 years. We found that Acadian Flycatcher abundance was initially greater in hardwood plantation habitat relative to bottomland hardwood forest along river levees but that abundance declined over time in both habitats. We plan for unmarked development to keep pace with advances in hierarchical modelling in ecology, including better handling of continuous-time data from camera trap and automated recording units and integrated models for multiple data streams.

Methods in Ecology & Evolution

Wetland selection by female Ring-Necked Ducks (Aythya collaris) in the Southern Atlantic Flyway

On the wintering grounds, wetland selection by waterfowl is influenced by spatiotemporal resource distribution. The ring-necked duck (Aythya collaris) winters in the southeastern United States where a disproportionate amount of Atlantic Flyway ring-necked duck harvest occurs. We quantified female ring-necked duck selection for wetland characteristics during and after the 2017-2018 and 2018-2019 waterfowl hunting seasons using discrete choice modeling under a Bayesian framework. Relative probability of selection was primarily influenced by characteristics at the local wetland scale. Relative probability of selection was higher for flooded agriculture and vegetated wetlands than open water and was positively influenced by wetland area during the winter. After the hunting season, the relative probability of selection decreased for flooded agriculture but increased for vegetated wetlands, and the effect of wetland area decreased in magnitude. We attribute changes in selection during and after the hunting season to dietary shifts related to migratory preparation, resource depletion, and reproductive pairing. Understanding the wetland characteristics that wintering waterfowl select, and the spatial scale at which selection occurs, is important for informing effective wetland management and waterfowl harvest practices.

Florida, Georgia

Spatial capture–recapture with random thinning for unidentified encounters

Spatial capture–recapture (SCR) models have increasingly been used as a basis for combining capture–recapture data types with variable levels of individual identity information to estimate population density and other demographic parameters. Recent examples are the unmarked SCR (or spatial count model), where no individual identities are available and spatial mark–resight (SMR) where individual identities are available for only a marked subset of the population. Currently lacking, though, is a model that allows unidentified samples to be combined with identified samples when there are no separate classes of “marked” and “unmarked” individuals and when the two sample types cannot be considered as arising from two independent observation models. This is a common scenario when using noninvasive sampling methods, for example, when analyzing data on identified and unidentified photographs or scats from the same sites. Here we describe a “random thinning” SCR model that utilizes encounters of both known and unknown identity samples using a natural mechanistic dependence between samples arising from a single observation model. Our model was fitted in a Bayesian framework using NIMBLE. We investigate the improvement in parameter estimates by including the unknown identity samples, which was notable (up to 79% more precise) in low‐density populations with a low rate of identified encounters. We then applied the random thinning SCR model to a noninvasive genetic sampling study of brown bear ( Ursus arctos ) density in Oriental Cantabrian Mountains (North Spain). Our model can improve density estimation for noninvasive sampling studies for low‐density populations with low rates of individual identification, by making use of available data that might otherwise be discarded.

Ecology & Evolution

Informing amphibian conservation efforts with abundance-based metapopulation models

Science-based management strategies are needed to halt or reverse the global decline of amphibians. In many cases, sound management requires reliable models built using monitoring data. Historically, monitoring and statistical modeling efforts have focused on estimating occupancy using detection–nondetection data. Spatial occupancy models are useful for studying colonization–extinction dynamics, but richer insights can be gained from estimating abundance and density-dependent demographic rates. We developed an integrated abundance-based metapopulation model of the processes contributing to spatiotemporal variation in patch population density. We fit our model to a combination of detection–nondetection and count data from a 14-yr study of a reintroduced metapopulation of federally threatened Chiricahua Leopard Frogs (Lithobates chiricahuensis). Pond-specific population growth rate was influenced by pond hydroperiod and frog density, such that permanent and semipermanent ponds with low densities of adult frogs experienced the highest annual population growth rates. Immigration rate declined as the distance among ponds increased. After reintroduction in 2003, metapopulation-level abundance increased and appeared to stabilize around 1300 adult frogs (95% CI = 1192–1471) by year 2015. Further, changes in metapopulation abundance were driven mostly by changes in abundance at a few ponds. These high-density populations, which would not have been identifiable with traditional occupancy-based metapopulation models, are likely especially important for species recovery in the area. Abundance-based metapopulation models can be widely applied to inform conservation efforts, by providing higher quality information needed to prioritize habitat patches for management and can be used to make more accurate predictions of metapopulation extinction risk.

Herpetologica

A statistical forecasting approach to metapopulation viability analysis

Conservation of at‐risk species is aided by reliable forecasts of the consequences of environmental change and management actions on population viability. Forecasts from conventional population viability analysis (PVA) are made using a two‐step procedure in which parameters are estimated, or elicited from expert opinion, and then plugged into a stochastic population model without accounting for parameter uncertainty. Recently developed statistical PVAs differ because forecasts are made conditional on models fitted to empirical data. The statistical forecasting approach allows for uncertainty about parameters, but it has rarely been applied in metapopulation contexts where spatially explicit inference is needed about colonization and extinction dynamics and other forms of stochasticity that influence metapopulation viability. We conducted a statistical metapopulation viability analysis (MPVA) using 11 yr of data on the federally threatened Chiricahua leopard frog ( Lithobates chiricahuensis ) to forecast responses to landscape heterogeneity, drought, environmental stochasticity, and management. We evaluated several future environmental scenarios and pond restoration options designed to reduce extinction risk. Forecasts over a 50‐yr time horizon indicated that metapopulation extinction risk was <4% for all scenarios, but uncertainty was high. Without pond restoration, extinction risk is forecasted to be 3.9% (95% CI 0–37%) by year 2066. Restoring six ponds by increasing their hydroperiod reduced extinction risk to <1% and greatly reduced uncertainty (95% CI 0–2%). Our results suggest that managers can mitigate the impacts of drought and environmental stochasticity on metapopulation viability by maintaining ponds that hold water throughout the year and keeping them free of invasive predators. Our study illustrates the utility of the spatially explicit statistical forecasting approach to MPVA in conservation planning efforts.

Ecological Applications

Spatial capture–recapture for categorically marked populations with an application to genetic capture–recapture

Recently introduced unmarked spatial capture–recapture (SCR), spatial mark–resight (SMR), and 2‐flank spatial partial identity models (SPIMs) extend the domain of SCR to populations or observation systems that do not always allow for individual identity to be determined with certainty. For example, some species do not have natural marks that can reliably produce individual identities from photographs, and some methods of observation produce partial identity samples as is the case with remote cameras that sometimes produce single‐flank photographs. Unmarked SCR, SMR, and SPIM share the feature that they probabilistically resolve the uncertainty in individual identity using the spatial location where samples were collected. Spatial location is informative of individual identity in spatially structured populations because a sample is more likely to have been produced by an individual living near the trap where it was recorded than an individual living further away from the trap. Further, the level of information about individual identity that a spatial location contains is related to two key ecological concepts, population density and home range size, which we quantify using a proposed Identity Diversity Index (IDI). We show that latent and partial identity SCR models produce imprecise and biased density estimates in many high IDI scenarios when data are sparse. We then extend the unmarked SCR model to incorporate categorical, partially identifying covariates, which reduce the level of uncertainty in individual identity, increasing the reliability and precision of density estimates, and allowing reliable density estimation in scenarios with higher IDI values and with more sparse data. We illustrate the performance of this “categorical SPIM” via simulations and by applying it to a black bear data set using microsatellite loci as categorical covariates, where we reproduce the full data set estimates with only slightly less precision using fewer loci than necessary for confident individual identification. We then discuss how the categorical SPIM can be applied to other wildlife sampling scenarios such as remote camera surveys, where natural or researcher‐applied partial marks can be observed in photographs. Finally, we discuss how the categorical SPIM can be added to SMR, 2‐flank SPIM, or other latent identity SCR models.

Ecosphere

Estimating population extinction thresholds with categorical classification trees for Louisiana black bears

Monitoring vulnerable species is critical for their conservation. Thresholds or tipping points are commonly used to indicate when populations become vulnerable to extinction and to trigger changes in conservation actions. However, quantitative methods to determine such thresholds have not been well explored. The Louisiana black bear ( Ursus americanus luteolus ) was removed from the list of threatened and endangered species under the U.S. Endangered Species Act in 2016 and our objectives were to determine the most appropriate parameters and thresholds for monitoring and management action. Capture mark recapture (CMR) data from 2006 to 2012 were used to estimate population parameters and variances. We used stochastic population simulations and conditional classification trees to identify demographic rates for monitoring that would be most indicative of heighted extinction risk. We then identified thresholds that would be reliable predictors of population viability. Conditional classification trees indicated that annual apparent survival rates for adult females averaged over 5 years ( ) was the best predictor of population persistence. Specifically, population persistence was estimated to be ≥95% over 100 years when , suggesting that this statistic can be used as threshold to trigger management intervention. Our evaluation produced monitoring protocols that reliably predicted population persistence and was cost-effective. We conclude that population projections and conditional classification trees can be valuable tools for identifying extinction thresholds used in monitoring programs.

PLoS ONE

Demographic rates and population viability of black bears in Louisiana

The Louisiana black bear ( Ursus americanus luteolus ) was reduced to a few small, fragmented, and isolated subpopulations in the Lower Mississippi Alluvial Valley by the mid-twentieth century resulting from loss and fragmentation of habitat. In 1992, the United States Fish and Wildlife Service (USFWS) granted the Louisiana black bear threatened status under the United States Endangered Species Act of 1973. Since that time, a recovery plan was developed, a reintroduced population was established, and habitat recovery has occurred. The Recovery Plan states that a minimum of 2 populations must be viable (i.e., persistence probabilities over 100 years >0.95), 1 in the Tensas River Basin and 1 in the Atchafalaya River Basin. Consequently, our objectives were to 1) estimate demographic rates of Louisiana black bear subpopulations, 2) develop data-driven stochastic population projection models, and 3) determine how different projection model assumptions affect population trajectories and predictions about long-term persistence. Our overall goal was to assess long-term persistence of the bear subpopulations in Louisiana, individually and as a whole. We collected data using varying combinations of non-invasive DNA sampling, live capture, winter den visits, and radio monitoring from 2002 to 2012 in the 4 areas currently supporting breeding subpopulations in Louisiana: Tensas River Basin (TRB), Upper Atchafalaya River Basin (UARB), Lower Atchafalaya River Basin (LARB), and a recently reintroduced population at the Three Rivers Complex (TRC). From 2002 to 2012, we radio monitored fates of 86 adult females within the TRB and 43 in the TRC. Mean estimates of annual adult survival for the TRB and TRC were 0.997 and 0.990, respectively, when unknown fates were assumed alive and 0.970 and 0.926 when unknown fates were assumed dead. From 2003 to 2013, we observed 130 cub litters from 74 females in the TRB, and 74 cub litters from 45 females in the TRC. During the same period, we observed 43 yearling litters for 33 females in the TRB and 21 yearling litters for 19 females in the TRC. The estimated number of cubs and number of yearlings produced per breeding adult female was 0.47 and 0.20, respectively, in the TRB and 0.32 and 0.18 in the TRC. On the basis of matrix projection models, asymptotic growth rates ranged from 1.053 to 1.078 for the TRB and from 1.005 to 1.062 for the TRC, depending on how we treated unresolved fates of adult females. Persistence probabilities estimated from stochastic population models based on telemetry data ranged from 0.997 to 0.998 for the TRC subpopulation depending on model assumptions and were >0.999 for the TRB regardless of model assumptions. We extracted DNA from hair collected at baited, barbed-wire enclosures in the TRB, UARB, and LARB to determine individual identities for capture-mark-recapture (CMR) analysis. We used those detection histories to estimate apparent survival ( φ ), per-capita recruitment ( f ), abundance ( N ), realized growth rate ( λ ), and long-term viability, based on Bayesian hierarchical modeling methods that allowed estimation of temporal process variance and parameter uncertainty. Based on 23,312 hair samples, annual N for females in the TRB ranged from 133 to 164 during 2006–2012, depending on year and how detection heterogeneity was modeled. Geometric mean of λ ranged from 0.996 to 1.002. In the UARB, we collected 11,643 hair samples from 2007 to 2012, from which estimates of N for females ranged from 23 to 43 during the study period, depending on detection heterogeneity model. The geometric mean of λ ranged from 1.038 to 1.059. Estimated N for females in LARB ranged from 69 to 96, and annual λ ranged from 0.80 to 1.11 based on 3,698 hair samples collected during 2010–2012, also depending on year and heterogeneity model. Probabilities of persistence over 100 years for the TRC and TRB based on stochastic matrix projection models that used vital rate estimates from telemetry data were >0.95 for all scenarios. Probability of persistence at the TRB and the UARB based on projection models that used vital rate estimates from CMR analyses ranged from 0.928 to 0.954 and from 0.906 to 0.959, respectively, depending on model assumptions. Data from the LARB were insufficient for a viability assessment. Thus, individual persistence probabilities for TRB and UARB did not meet the strict definition of viability (i.e., >0.95) under some model assumptions. However, the joint probability of bears persisting either in the TRB or UARB was >0.993 assuming individual population dynamics were independent and was >0.958 assuming dynamics were perfectly correlated. Furthermore, including the TRC increased the joint probability of bears persisting somewhere in the TRB, UARB, or TRC to >0.999 based on the most pessimistic individual persistence estimates from those subpopulations. Therefore, if the intent of specifying that 2 subpopulations should be viable was to ensure the persistence of Louisiana black bears somewhere within its historical range, then the viability threshold was met. © 2016 The Wildlife Society.

Louisiana

Survival estimates for reintroduced populations of the Chiricahua Leopard Frog ( Lithobates chiricahuensis )

Global amphibian declines have been attributed to a number of factors including disease, invasive species, habitat degradation, and climate change. Reintroduction is one management action that is commonly used with the goal of recovering imperiled species. The success of reintroductions varies widely, and evaluating their efficacy requires estimates of population viability metrics, such as underlying vital rates and trends in abundance. Although rarely quantified, assessing vital rates for recovering populations provides a more mechanistic understanding of population growth than numerical trends in population occupancy or abundance. We used three years of capture-mark-recapture data from three breeding ponds and a Cormack-Jolly-Seber model to estimate annual apparent survival for reintroduced populations of the federally threatened Chiricahua Leopard Frog ( Lithobates chiricahuensis ) at the Buenos Aires National Wildlife Refuge (BANWR), in the Altar Valley, Arizona, USA. To place our results in context, we also compiled published survival estimates for other ranids. Average apparent survival of Chiricahua Leopard Frogs at BANWR was 0.27 (95% CI [0.07, 0.74]) and average individual capture probability was 0.02 (95% CI [0, 0.05]). Our apparent survival estimate for Chiricahua Leopard Frogs is lower than for most other ranids and is not consistent with recent research that showed metapopulation viability in the Altar Valley is high. We suggest that low apparent survival may be indicative of high emigration rates. We recommend that future research should estimate emigration rates so that actual, rather than apparent, survival can be quantified to improve population viability assessments of threatened species following reintroduction efforts.

Copeia

Spatial occupancy models for predicting metapopulation dynamics and viability following reintroduction

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

Arizona, New Mexico

Estimating landscape resistance to dispersal

Dispersal is an inherently spatial process that can be affected by habitat conditions in sites encountered by dispersers. Understanding landscape resistance to dispersal is important in connectivity studies and reserve design, but most existing methods use resistance functions with cost parameters that are subjectively chosen by the investigator. We develop an analytic approach allowing for direct estimation of resistance parameters that folds least cost path methods typically used in simulation approaches into a formal statistical model of dispersal distributions. The core of our model is a frequency distribution of dispersal distances expressed as least cost distance rather than Euclidean distance, and which includes terms for feature-specific costs to dispersal and sex (or other traits) of the disperser. The model requires only origin and settlement locations for multiple individuals, such as might be obtained from mark&ndash;recapture studies or parentage analyses, and maps of the relevant habitat features. To evaluate whether the model can estimate parameters correctly, we fit our model to data from simulated dispersers in three kinds of landscapes (in which resistance of environmental variables was categorical, continuous with a patchy configuration, or continuous in a trend pattern). We found maximum likelihood estimators of resistance and individual trait parameters to be approximately unbiased with moderate sample sizes. We applied the model to a small grizzly bear dataset to demonstrate how this approach could be used when the primary interest is in the prediction of costs and found that estimates were consistent with expectations based on bear ecology. Our method has important practical applications for testing hypotheses about dispersal ecology and can be used to inform connectivity planning efforts, via the resistance estimates and confidence intervals, which can be used to create a data-driven resistance surface.

Landscape Ecology

Integrating resource selection information with spatial capture--recapture

1. Understanding space usage and resource selection is a primary focus of many studies of animal populations. Usually, such studies are based on location data obtained from telemetry, and resource selection functions (RSFs) are used for inference. Another important focus of wildlife research is estimation and modeling population size and density. Recently developed spatial capture–recapture (SCR) models accomplish this objective using individual encounter history data with auxiliary spatial information on location of capture. SCR models include encounter probability functions that are intuitively related to RSFs, but to date, no one has extended SCR models to allow for explicit inference about space usage and resource selection. 2. In this paper we develop the first statistical framework for jointly modeling space usage, resource selection, and population density by integrating SCR data, such as from camera traps, mist-nets, or conventional catch traps, with resource selection data from telemetered individuals. We provide a framework for estimation based on marginal likelihood, wherein we estimate simultaneously the parameters of the SCR and RSF models. 3. Our method leads to increases in precision for estimating parameters of ordinary SCR models. Importantly, we also find that SCR models alone can estimate parameters of RSFs and, as such, SCR methods can be used as the sole source for studying space-usage; however, precision will be higher when telemetry data are available. 4. Finally, we find that SCR models using standard symmetric and stationary encounter probability models may not fully explain variation in encounter probability due to space usage, and therefore produce biased estimates of density when animal space usage is related to resource selection. Consequently, it is important that space usage be taken into consideration, if possible, in studies focused on estimating density using capture–recapture methods.

Methods in Ecology and Evolution

Spatially explicit models for inference about density in unmarked or partially marked populations

Recently developed spatial capture–recapture (SCR) models represent a major advance over traditional capture–recapture (CR) models because they yield explicit estimates of animal density instead of population size within an unknown area. Furthermore, unlike nonspatial CR methods, SCR models account for heterogeneity in capture probability arising from the juxtaposition of animal activity centers and sample locations. Although the utility of SCR methods is gaining recognition, the requirement that all individuals can be uniquely identified excludes their use in many contexts. In this paper, we develop models for situations in which individual recognition is not possible, thereby allowing SCR concepts to be applied in studies of unmarked or partially marked populations. The data required for our model are spatially referenced counts made on one or more sample occasions at a collection of closely spaced sample units such that individuals can be encountered at multiple locations. Our approach includes a spatial point process for the animal activity centers and uses the spatial correlation in counts as information about the number and location of the activity centers. Camera-traps, hair snares, track plates, sound recordings, and even point counts can yield spatially correlated count data, and thus our model is widely applicable. A simulation study demonstrated that while the posterior mean exhibits frequentist bias on the order of 5–10% in small samples, the posterior mode is an accurate point estimator as long as adequate spatial correlation is present. Marking a subset of the population substantially increases posterior precision and is recommended whenever possible. We applied our model to avian point count data collected on an unmarked population of the northern parula (Parula americana) and obtained a density estimate (posterior mode) of 0.38 (95% CI: 0.19–1.64) birds/ha. Our paper challenges sampling and analytical conventions in ecology by demonstrating that neither spatial independence nor individual recognition is needed to estimate population density—rather, spatial dependence can be informative about individual distribution and density.

Annals of Applied Statistics

Spatial capture-recapture models for jointly estimating population density and landscape connectivity

Population size and landscape connectivity are key determinants of population viability, yet no methods exist for simultaneously estimating density and connectivity parameters. Recently developed spatial capture–recapture (SCR) models provide a framework for estimating density of animal populations but thus far have not been used to study connectivity. Rather, all applications of SCR models have used encounter probability models based on the Euclidean distance between traps and animal activity centers, which implies that home ranges are stationary, symmetric, and unaffected by landscape structure. In this paper we devise encounter probability models based on “ecological distance,” i.e., the least-cost path between traps and activity centers, which is a function of both Euclidean distance and animal movement behavior in resistant landscapes. We integrate least-cost path models into a likelihood-based estimation scheme for spatial capture–recapture models in order to estimate population density and parameters of the least-cost encounter probability model. Therefore, it is possible to make explicit inferences about animal density, distribution, and landscape connectivity as it relates to animal movement from standard capture–recapture data. Furthermore, a simulation study demonstrated that ignoring landscape connectivity can result in negatively biased density estimators under the naive SCR model.

Ecology

Spatial capture-recapture

Spatial Capture-Recapture provides a revolutionary extension of traditional capture-recapture methods for studying animal populations using data from live trapping, camera trapping, DNA sampling, acoustic sampling, and related field methods. This book is a conceptual and methodological synthesis of spatial capture-recapture modeling. As a comprehensive how-to manual, this reference contains detailed examples of a wide range of relevant spatial capture-recapture models for inference about population size and spatial and temporal variation in demographic parameters. Practicing field biologists studying animal populations will find this book to be a useful resource, as will graduate students and professionals in ecology, conservation biology, and fisheries and wildlife management.

Book

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

Hierarchical distance-sampling models to estimate population size and habitat-specific abundance of an island endemic

Population size and habitat-specific abundance estimates are essential for conservation management. A major impediment to obtaining such estimates is that few statistical models are able to simultaneously account for both spatial variation in abundance and heterogeneity in detection probability, and still be amenable to large-scale applications. The hierarchical distance-sampling model of J. A. Royle, D. K. Dawson, and S. Bates provides a practical solution. Here, we extend this model to estimate habitat-specific abundance and rangewide population size of a bird species of management concern, the Island Scrub-Jay ( Aphelocoma insularis ), which occurs solely on Santa Cruz Island, California, USA. We surveyed 307 randomly selected, 300 m diameter, point locations throughout the 250-km 2 island during October 2008 and April 2009. Population size was estimated to be 2267 (95% CI 1613-3007) and 1705 (1212-2369) during the fall and spring respectively, considerably lower than a previously published but statistically problematic estimate of 12 500. This large discrepancy emphasizes the importance of proper survey design and analysis for obtaining reliable information for management decisions. Jays were most abundant in low-elevation chaparral habitat; the detection function depended primarily on the percent cover of chaparral and forest within count circles. Vegetation change on the island has been dramatic in recent decades, due to release from herbivory following the eradication of feral sheep ( Ovis aries ) from the majority of the island in the mid-1980s. We applied best-fit fall and spring models of habitat-specific jay abundance to a vegetation map from 1985, and estimated the population size of A. insularis was 1400-1500 at that time. The 20-30% increase in the jay population suggests that the species has benefited from the recovery of native vegetation since sheep removal. Nevertheless, this jay's tiny range and small population size make it vulnerable to natural disasters and to habitat alteration related to climate change. Our results demonstrate that hierarchical distance-sampling models hold promise for estimating population size and spatial density variation at large scales. Our statistical methods have been incorporated into the R package unmarked to facilitate their use by animal ecologists, and we provide annotated code in the Supplement.

California

Unmarked: An R package for fitting hierarchical models of wildlife occurrence and abundance

Ecological research uses data collection techniques that are prone to substantial and unique types of measurement error to address scientific questions about species abundance and distribution. These data collection schemes include a number of survey methods in which unmarked individuals are counted, or determined to be present, at spatially- referenced sites. Examples include site occupancy sampling, repeated counts, distance sampling, removal sampling, and double observer sampling. To appropriately analyze these data, hierarchical models have been developed to separately model explanatory variables of both a latent abundance or occurrence process and a conditional detection process. Because these models have a straightforward interpretation paralleling mechanisms under which the data arose, they have recently gained immense popularity. The common hierarchical structure of these models is well-suited for a unified modeling interface. The R package unmarked provides such a unified modeling framework, including tools for data exploration, model fitting, model criticism, post-hoc analysis, and model comparison.

Journal of Statistical Software