Sharing land via keystone structure: Retaining naturally regenerated trees may efficiently benefit birds in plantations.

Meeting food/wood demands with increasing human population and per-capita consumption is a pressing conservation issue, and is often framed as a choice between land sparing and land sharing. Although most empirical studies comparing the efficacy of land sparing and sharing supported land sparing, land sharing may be more efficient if its performance is tested by rigorous experimental design and habitat structures providing crucial resources for various species-keystone structures-are clearly involved. We launched a manipulative experiment to retain naturally regenerated broad-leaved trees when harvesting conifer plantations in central Hokkaido, northern Japan. We surveyed birds in harvested treatments, unharvested plantation controls, and natural forest references 1-year before the harvest and for three consecutive postharvest years. We developed a hierarchical community model separating abundance and space use (territorial proportion overlapping treatment plots) subject to imperfect detection to assess population consequences of retention harvesting. Application of the model to our data showed that retaining some broad-leaved trees increased the total abundance of forest birds over the harvest rotation cycle. Specifically, a preharvest survey showed that the amount of broad-leaved trees increased forest bird abundance in a concave manner (i.e., in the form of diminishing returns). After harvesting, a small amount of retained broad-leaved trees mitigated negative harvesting impacts on abundance, although retention harvesting reduced the space use. Nevertheless, positive retention effects on the postharvest bird density as the product of abundance and space use exhibited a concave form. Thus, small profit reductions were shown to yield large increases in forest bird abundance. The difference in bird abundance between clearcutting and low amounts of broad-leaved tree retention increased slightly from the first to second postharvesting years. We conclude that retaining a small amount of broad-leaved trees may be a cost-effective on-site conservation approach for the management of conifer plantations. The retention of 20-30 broad-leaved trees per ha may be sufficient to maintain higher forest bird abundance than clearcutting over the rotation cycle. Retention approaches can be incorporated into management systems using certification schemes and best management practices. Developing an awareness of the roles and values of naturally regenerated trees is needed to diversify plantations.

Spatial proximity moderates genotype uncertainty in genetic tagging studies.

Accelerating declines of an increasing number of animal populations worldwide necessitate methods to reliably and efficiently estimate demographic parameters such as population density and trajectory. Standard methods for estimating demographic parameters from noninvasive genetic samples are inefficient because lower-quality samples cannot be used, and they assume individuals are identified without error. We introduce the genotype spatial partial identity model (gSPIM), which integrates a genetic classification model with a spatial population model to combine both spatial and genetic information, thus reducing genotype uncertainty and increasing the precision of demographic parameter estimates. We apply this model to data from a study of fishers (Pekania pennanti) in which 37% of hair samples were originally discarded because of uncertainty in individual identity. The gSPIM density estimate using all collected samples was 25% more precise than the original density estimate, and the model identified and corrected three errors in the original individual identity assignments. A simulation study demonstrated that our model increased the accuracy and precision of density estimates 63 and 42%, respectively, using three replicated assignments (e.g., PCRs for microsatellites) per genetic sample. Further, the simulations showed that the gSPIM model parameters are identifiable with only one replicated assignment per sample and that accuracy and precision are relatively insensitive to the number of replicated assignments for high-quality samples. Current genotyping protocols devote the majority of resources to replicating and confirming high-quality samples, but when using the gSPIM, genotyping protocols could be more efficient by devoting more resources to low-quality samples.

Migratory behavior and winter geography drive differential range shifts of eastern birds in response to recent climate change.

Over the past half century, migratory birds in North America have shown divergent population trends relative to resident species, with the former declining rapidly and the latter increasing. The role that climate change has played in these observed trends is not well understood, despite significant warming over this period. We used 43 y of monitoring data to fit dynamic species distribution models and quantify the rate of latitudinal range shifts in 32 species of birds native to eastern North America. Since the early 1970s, species that remain in North America throughout the year, including both resident and migratory species, appear to have responded to climate change through both colonization of suitable area at the northern leading edge of their breeding distributions and adaption in place at the southern trailing edges. Neotropical migrants, in contrast, have shown the opposite pattern: contraction at their southern trailing edges and no measurable shifts in their northern leading edges. As a result, the latitudinal distributions of temperate-wintering species have increased while the latitudinal distributions of neotropical migrants have decreased. These results raise important questions about the mechanisms that determine range boundaries of neotropical migrants and suggest that these species may be particularly vulnerable to future climate change. Our results highlight the potential importance of climate change during the nonbreeding season in constraining the response of migratory species to temperature changes at both the trailing and leading edges of their breeding distributions. Future research on the interactions between breeding and nonbreeding climate change is urgently needed.

Estimating and forecasting spatial population dynamics of apex predators using transnational genetic monitoring.

The ongoing recovery of terrestrial large carnivores in North America and Europe is accompanied by intense controversy. On the one hand, reestablishment of large carnivores entails a recovery of their most important ecological role, predation. On the other hand, societies are struggling to relearn how to live with apex predators that kill livestock, compete for game species, and occasionally injure or kill people. Those responsible for managing these species and mitigating conflict often lack fundamental information due to a long-standing challenge in ecology: How do we draw robust population-level inferences for elusive animals spread over immense areas? Here we showcase the application of an effective tool for spatially explicit tracking and forecasting of wildlife population dynamics at scales that are relevant to management and conservation. We analyzed the world's largest dataset on carnivores comprising more than 35,000 noninvasively obtained DNA samples from over 6,000 individual brown bears (Ursus arctos), gray wolves (Canis lupus), and wolverines (Gulo gulo). Our analyses took into account that not all individuals are detected and, even if detected, their fates are not always known. We show unequivocal quantitative evidence of large carnivore recovery in northern Europe, juxtaposed with the finding that humans are the single-most important factor driving the dynamics of these apex predators. We present maps and forecasts of the spatiotemporal dynamics of large carnivore populations, transcending national boundaries and management regimes.

Leveraging community science data for population assessments during a pandemic.

The COVID-19 pandemic has disrupted field research programs, making conservation and management decision-making more challenging. However, it may be possible to conduct population assessments using integrated models that combine community science data with existing data from structured surveys. We developed a space-time integrated model to characterize spatial and temporal variability in population distribution. We fit our integrated model to 10 years of eBird (2010-2020) and 9 years of aerial survey (2010-2019) Mottled Duck count data to forecast 2020 population size along the western Gulf Coast of Texas and Louisiana. Estimates of Mottled Duck abundance were similar in magnitude to estimates calculated using previous methods but were more precise and showed evidence of a declining population. The spatial distribution for Mottled Ducks each year was characterized by several concentrations of relatively high abundance, although the location of these abundance "hotspots" varied over time. Expected abundance was higher for areas with a higher proportion of area covered by marsh habitat. By leveraging large-scale community science data, we were able to conduct a population assessment despite the disruption in structured surveys caused by the pandemic. As participation in community science platforms continues to increase, we anticipate modeling frameworks, like the integrated model we developed here, will become increasingly useful for informing conservation and management decision-making.

Evaluation of a two-season banding program to estimate and model migratory bird survival.

The management of North American waterfowl is predicated on long-term, continental-scale banding implemented prior to the hunting season (i.e., July-September) and subsequent reporting of bands recovered by hunters. However, single-season banding and encounter operations have a number of characteristics that limit their application to estimating demographic rates and evaluating hypothesized limiting factors throughout the annual cycle. We designed and implemented a two-season banding program for American Black Ducks (Anas rubripes), Mallards (A. platyrhynchos), and hybrids in eastern North America to evaluate potential application to annual life cycle conservation and sport harvest management. We assessed model fit and compared estimates of annual survival among data types (i.e., pre-hunting season only [July-September], post-hunting season only [January-March], and two-season [pre- and post-hunting season]) to evaluate model assumptions and potential application to population modeling and management. There was generally high agreement between estimates of annual survival derived using two-season and pre-season only data for all age and sex cohorts. Estimates of annual survival derived from post-season banding data only were consistently higher for adult females and juveniles of both sexes. We found patterns of seasonal survival varied by species, age, and to a lesser extent, sex. Hunter recovered birds exhibited similar spatial distributions regardless of banding season suggesting banded samples were from the same population. In contrast, goodness-of-fit tests suggest this assumption was statistically violated in some regions and years. We conclude that estimates of seasonal and annual survival for Black Ducks and Mallards based on the two-season banding program are valid and accurate based on model fit statistics, similarity in survival estimates across data and models, and similarities in the distribution of recoveries. The two-season program provides greater precision and insight into the survival process and will improve the ability of researchers and managers to test competing hypotheses regarding population regulation resulting in more effective management.

Genetic tagging in the Anthropocene: scaling ecology from alleles to ecosystems.

The Anthropocene is an era of marked human impact on the world. Quantifying these impacts has become central to understanding the dynamics of coupled human-natural systems, resource-dependent livelihoods, and biodiversity conservation. Ecologists are facing growing pressure to quantify the size, distribution, and trajectory of wild populations in a cost-effective and socially acceptable manner. Genetic tagging, combined with modern computational and genetic analyses, is an under-utilized tool to meet this demand, especially for wide-ranging, elusive, sensitive, and low-density species. Genetic tagging studies are now revealing unprecedented insight into the mechanisms that control the density, trajectory, connectivity, and patterns of human-wildlife interaction for populations over vast spatial extents. Here, we outline the application of, and ecological inferences from, new analytical techniques applied to genetically tagged individuals, contrast this approach with conventional methods, and describe how genetic tagging can be better applied to address outstanding questions in ecology. We provide example analyses using a long-term genetic tagging dataset of grizzly bears in the Canadian Rockies. The genetic tagging toolbox is a powerful and overlooked ensemble that ecologists and conservation biologists can leverage to generate evidence and meet the challenges of the Anthropocene.

Reserve design to optimize functional connectivity and animal density.

Ecological distance-based spatial capture-recapture models (SCR) are a promising approach for simultaneously estimating animal density and connectivity, both of which affect spatial population processes and ultimately species persistence. We explored how SCR models can be integrated into reserve-design frameworks that explicitly acknowledge both the spatial distribution of individuals and their space use resulting from landscape structure. We formulated the design of wildlife reserves as a budget-constrained optimization problem and conducted a simulation to explore 3 different SCR-informed optimization objectives that prioritized different conservation goals by maximizing the number of protected individuals, reserve connectivity, and density-weighted connectivity. We also studied the effect on our 3 objectives of enforcing that the space-use requirements of individuals be met by the reserve for individuals to be considered conserved (referred to as home-range constraints). Maximizing local population density resulted in fragmented reserves that would likely not aid long-term population persistence, and maximizing the connectivity objective yielded reserves that protected the fewest individuals. However, maximizing density-weighted connectivity or preemptively imposing home-range constraints on reserve design yielded reserves of largely spatially compact sets of parcels covering high-density areas in the landscape with high functional connectivity between them. Our results quantify the extent to which reserve design is constrained by individual home-range requirements and highlight that accounting for individual space use in the objective and constraints can help in the design of reserves that balance abundance and connectivity in a biologically relevant manner.

Diseño de Reservas para Optimizar la Conectividad Funcional y la Densidad Animal Resumen Los modelos de captura-recaptura espacial (CRE) basados en distancias ecológicas son un método prometedor para estimar la densidad animal y la conectividad, las cuales afectan los procesos poblacionales espaciales y, en última instancia, la persistencia de las especies. Exploramos cómo se puede integrar a los modelos CRE en los marcos de diseño de reserva que explícitamente reconocen tanto la distribución espacial de los individuos como su uso del espacio resultante de la estructura del paisaje. Formulamos el diseño de reservas de vida silvestre como un problema de optimización de presupuesto limitado y realizamos una simulación para explorar 3 diferentes objetivos de optimización informados por CRE que priorizaron diferentes metas de conservación mediante la maximización del número de individuos protegidos; la conectividad de la reserva y la conectividad ponderada por la densidad. También estudiamos el efecto sobre nuestros objetivos de hacer que los requerimientos individuales de uso de espacio fuesen satisfechos por la reserva de manera que se pudiese considerar que los individuos estaban protegidos (referidos como restricciones de rango de hogar). La maximización de la densidad de la población local resultó en reservas fragmentadas que probablemente no contribuyan a la persistencia de la población a largo plazo, mientras que la maximización de la conectividad produjo reservas que protegían al menor número de individuos. Sin embargo, la maximización de la conectividad ponderada por la densidad o la imposición preventiva de restricciones de rango de hogar en el diseño de reservas produjo reservas compuestas por conjuntos de parcelas mayormente compactas espacialmente que cubrían áreas de densidad alta en el paisaje con alta conectividad funcional entre ellas. Nuestros resultados cuantifican la extensión a la cual el diseño de reservas esta limitado por los requerimientos de rango de hogar individuales y resaltan que la consideración del uso de espacio individual en el objetivo y limitaciones puede ayudar al diseño de reservas que equilibren la abundancia y la conectividad de manera biológicamente relevante.

Eco-evolutionary rescue promotes host-pathogen coexistence.

Emerging infectious pathogens are responsible for some of the most severe host mass mortality events in wild populations. Yet, effective pathogen control strategies are notoriously difficult to identify, in part because quantifying and forecasting pathogen spread and disease dynamics is challenging. Following an outbreak, hosts must cope with the presence of the pathogen, leading to host-pathogen coexistence or extirpation. Despite decades of research, little is known about host-pathogen coexistence post-outbreak when low host abundances and cryptic species make these interactions difficult to study. Using a novel disease-structured N-mixture model, we evaluate empirical support for three host-pathogen coexistence hypotheses (source-sink, eco-evolutionary rescue, and spatial variation in pathogen transmission) in a Neotropical amphibian community decimated by Batrachochytrium dendrobatidis (Bd) in 2004. During 2010-2014, we surveyed amphibians in Parque Nacional G. D. Omar Torríjos Herrera, Coclé Province, El Copé, Panama. We found that the primary driver of host-pathogen coexistence was eco-evolutionary rescue, as evidenced by similar amphibian survival and recruitment rates between infected and uninfected hosts. Average apparent monthly survival rates of uninfected and infected hosts were both close to 96%, and the expected number of uninfected and infected hosts recruited (via immigration/reproduction) was less than one host per disease state per 20-m site. The secondary driver of host-pathogen coexistence was spatial variation in pathogen transmission as we found that transmission was highest in areas of low abundance but there was no support for the source-sink hypothesis. Our results indicate that changes in the host community (i.e., through genetic or species composition) can reduce the impacts of emerging infectious disease post-outbreak. Our disease-structured N-mixture model represents a valuable advancement for conservation managers trying to understand underlying host-pathogen interactions and provides new opportunities to study disease dynamics in remnant host populations decimated by virulent pathogens.

An integrated population model for bird monitoring in North America.

Integrated population models (IPMs) provide a unified framework for simultaneously analyzing data sets of different types to estimate vital rates, population size, and dynamics; assess contributions of demographic parameters to population changes; and assess population viability. Strengths of an IPM include the ability to estimate latent parameters and improve the precision of parameter estimates. We present a hierarchical IPM that combines two broad-scale avian monitoring data sets: count data from the North American Breeding Bird Survey (BBS) and capture-recapture data from the Monitoring Avian Productivity and Survivorship (MAPS) program. These data sets are characterized by large numbers of sample sites and observers, factors capable of inducing error in the sampling and observation processes. The IPM integrates the data sets by modeling the population abundance as a first-order autoregressive function of the previous year's population abundance and vital rates. BBS counts were modeled as a log-linear function of the annual index of population abundance, observation effects (observer identity and first survey year), and overdispersion. Vital rates modeled included adult apparent survival, estimated from a transient Cormack-Jolly-Seber model using MAPS data, and recruitment (surviving hatched birds from the previous season + dispersing adults) estimated as a latent parameter. An assessment of the IPM demonstrated it could recover true parameter values from 200 simulated data sets. The IPM was applied to data sets (1992-2008) of two bird species, Gray Catbird (Dumetella carolinensis) and Wood Thrush (Hylocichla mustelina) in the New England/Mid-Atlantic coastal Bird Conservation Region of the United States. The Gray Catbird population was relatively stable (trend +0.4% per yr), while the Wood Thrush population nearly halved (trend -4.5% per yr) over the 17-yr study period. IPM estimates of population growth rates, adult survival, and detection and residency probabilities were similar and as precise as estimates from the stand-alone BBS and CJS models. A benefit of using the IPM was its ability to estimate the latent recruitment parameter. Annual growth rates for both species correlated more with recruitment than survival, and the relationship for Wood Thrush was stronger than for Gray Catbird. The IPM's unified modeling framework facilitates integration of these important data sets.

Population size and stopover duration estimation using mark-resight data and Bayesian analysis of a superpopulation model.

We present a novel formulation of a mark-recapture-resight model that allows estimation of population size, stopover duration, and arrival and departure schedules at migration areas. Estimation is based on encounter histories of uniquely marked individuals and relative counts of marked and unmarked animals. We use a Bayesian analysis of a state-space formulation of the Jolly-Seber mark-recapture model, integrated with a binomial model for counts of unmarked animals, to derive estimates of population size and arrival and departure probabilities. We also provide a novel estimator for stopover duration that is derived from the latent state variable representing the interim between arrival and departure in the state-space model. We conduct a simulation study of field sampling protocols to understand the impact of superpopulation size, proportion marked, and number of animals sampled on bias and precision of estimates. Simulation results indicate that relative bias of estimates of the proportion of the population with marks was low for all sampling scenarios and never exceeded 2%. Our approach does not require enumeration of all unmarked animals detected or direct knowledge of the number of marked animals in the population at the time of the study. This provides flexibility and potential application in a variety of sampling situations (e.g., migratory birds, breeding seabirds, sea turtles, fish, pinnipeds, etc.). Application of the methods is demonstrated with data from a study of migratory sandpipers.

Estimating population density and connectivity of American mink using spatial capture-recapture.

Estimating the abundance or density of populations is fundamental to the conservation and management of species, and as landscapes become more fragmented, maintaining landscape connectivity has become one of the most important challenges for biodiversity conservation. Yet these two issues have never been formally integrated together in a model that simultaneously models abundance while accounting for connectivity of a landscape. We demonstrate an application of using capture-recapture to develop a model of animal density using a least-cost path model for individual encounter probability that accounts for non-Euclidean connectivity in a highly structured network. We utilized scat detection dogs (Canis lupus familiaris) as a means of collecting non-invasive genetic samples of American mink (Neovison vison) individuals and used spatial capture-recapture models (SCR) to gain inferences about mink population density and connectivity. Density of mink was not constant across the landscape, but rather increased with increasing distance from city, town, or village centers, and mink activity was associated with water. The SCR model allowed us to estimate the density and spatial distribution of individuals across a 388 km² area. The model was used to investigate patterns of space usage and to evaluate covariate effects on encounter probabilities, including differences between sexes. This study provides an application of capture-recapture models based on ecological distance, allowing us to directly estimate landscape connectivity. This approach should be widely applicable to provide simultaneous direct estimates of density, space usage, and landscape connectivity for many species.

An open-population hierarchical distance sampling model.

Modeling population dynamics while accounting for imperfect detection is essential to monitoring programs. Distance sampling allows estimating population size while accounting for imperfect detection, but existing methods do not allow for estimation of demographic parameters. We develop a model that uses temporal correlation in abundance arising from underlying population dynamics to estimate demographic parameters from repeated distance sampling surveys. Using a simulation study motivated by designing a monitoring program for Island Scrub-Jays (Aphelocoma insularis), we investigated the power of this model to detect population trends. We generated temporally autocorrelated abundance and distance sampling data over six surveys, using population rates of change of 0.95 and 0.90. We fit the data generating Markovian model and a mis-specified model with a log-linear time effect on abundance, and derived post hoc trend estimates from a model estimating abundance for each survey separately. We performed these analyses for varying numbers of survey points. Power to detect population changes was consistently greater under the Markov model than under the alternatives, particularly for reduced numbers of survey points. The model can readily be extended to more complex demographic processes than considered in our simulations. This novel framework can be widely adopted for wildlife population monitoring.

Modeling structured population dynamics using data from unmarked individuals.

The study of population dynamics requires unbiased, precise estimates of abundance and vital rates that account for the demographic structure inherent in all wildlife and plant populations. Traditionally, these estimates have only been available through approaches that rely on intensive mark-recapture data. We extended recently developed N-mixture models to demonstrate how demographic parameters and abundance can be estimated for structured populations using only stage-structured count data. Our modeling framework can be used to make reliable inferences on abundance as well as recruitment, immigration, stage-specific survival, and detection rates during sampling. We present a range of simulations to illustrate the data requirements, including the number of years and locations necessary for accurate and precise parameter estimates. We apply our modeling framework to a population of northern dusky salamanders (Desmognathus fuscus) in the mid-Atlantic region (USA) and find that the population is unexpectedly declining. Our approach represents a valuable advance in the estimation of population dynamics using multistate data from unmarked individuals and should additionally be useful in the development of integrated models that combine data from intensive (e.g., mark-recapture) and extensive (e.g., counts) data sources.

Integrated distance sampling models for simple point counts.

Point counts (PCs) are widely used in biodiversity surveys but, despite numerous advantages, simple PCs suffer from several problems: detectability, and therefore abundance, is unknown; systematic spatiotemporal variation in detectability yields biased inferences, and unknown survey area prevents formal density estimation and scaling-up to the landscape level. We introduce integrated distance sampling (IDS) models that combine distance sampling (DS) with simple PC or detection/nondetection (DND) data to capitalize on the strengths and mitigate the weaknesses of each data type. Key to IDS models is the view of simple PC and DND data as aggregations of latent DS surveys that observe the same underlying density process. This enables the estimation of separate detection functions, along with distinct covariate effects, for all data types. Additional information from repeat or time-removal surveys, or variable survey duration, enables the separate estimation of the availability and perceptibility components of detectability with DS and PC data. IDS models reconcile spatial and temporal mismatches among data sets and solve the above-mentioned problems of simple PC and DND data. To fit IDS models, we provide JAGS code and the new "IDS()" function in the R package unmarked. Extant citizen-science data generally lack the information necessary to adjust for detection biases, but IDS models address this shortcoming, thus greatly extending the utility and reach of these data. In addition, they enable formal density estimation in hybrid designs, which efficiently combine DS with distance-free, point-based PC or DND surveys. We believe that IDS models have considerable scope in ecology, management, and monitoring.

Current approaches using genetic distances produce poor estimates of landscape resistance to interindividual dispersal.

Landscape resistance reflects how difficult it is for genes to move across an area with particular attributes (e.g. land cover, slope). An increasingly popular approach to estimate resistance uses Mantel and partial Mantel tests or causal modelling to relate observed genetic distances to effective distances under alternative sets of resistance parameters. Relatively few alternative sets of resistance parameters are tested, leading to relatively poor coverage of the parameter space. Although this approach does not explicitly model key stochastic processes of gene flow, including mating, dispersal, drift and inheritance, bias and precision of the resulting resistance parameters have not been assessed. We formally describe the most commonly used model as a set of equations and provide a formal approach for estimating resistance parameters. Our optimization finds the maximum Mantel r when an optimum exists and identifies the same resistance values as current approaches when the alternatives evaluated are near the optimum. Unfortunately, even where an optimum existed, estimates from the most commonly used model were imprecise and were typically much smaller than the simulated true resistance to dispersal. Causal modelling using Mantel significance tests also typically failed to support the true resistance to dispersal values. For a large range of scenarios, current approaches using a simple correlational model between genetic and effective distances do not yield accurate estimates of resistance to dispersal. We suggest that analysts consider the processes important to gene flow for their study species, model those processes explicitly and evaluate the quality of estimates resulting from their model.

Markov models for community dynamics allowing for observation error.

Markov models are dynamic models that characterize transitions among discrete ecological states with transition probability matrices. Such models are widely used to infer community dynamics of sessile organisms because transition probabilities (the elements of transition probability matrices) can be estimated with time series data from "grid sampling," where species occupancy states are assessed at multiple fixed points in a quadrat or transect. These estimates, however, are known to be biased when resampling error exists. In this study, we used the perspective of multistate dynamic occupancy models to develop a new Markov model that is structured hierarchically such that transitions among occupancy states and observation processes are considered explicitly at each fixed point. We show that, by adopting a hierarchical Bayesian approach, our model provides estimates for transition probabilities that are robust to sampling error. We also show that error rate may be estimated without additional data obtained from rapid repeated sampling. Considerations for the analysis for the application to real data set and potential extensions of the proposed model are discussed.

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.

Density-habitat relationships of white-tailed deer (Odocoileus virginianus) in Finland.

In heterogeneous landscapes, resource selection constitutes a crucial link between landscape and population-level processes such as density. We conducted a non-invasive genetic study of white-tailed deer in southern Finland in 2016 and 2017 using fecal DNA samples to understand factors influencing white-tailed deer density and space use in late summer prior to the hunting season. We estimated deer density as a function of landcover types using a spatial capture-recapture (SCR) model with individual identities established using microsatellite markers. The study revealed second-order habitat selection with highest deer densities in fields and mixed forest, and third-order habitat selection (detection probability) for transitional woodlands (clear-cuts) and closeness to fields. Including landscape heterogeneity improved model fit and increased inferred total density compared with models assuming a homogenous landscape. Our findings underline the importance of including habitat covariates when estimating density and exemplifies that resource selection can be studied using non-invasive methods.