That's not the Mona Lisa! How to interpret spatial capture-recapture density surface estimates.

Spatial capture-recapture methods are often used to produce density surfaces, and these surfaces are often misinterpreted. In particular, spatial change in density is confused with spatial change in uncertainty about density. We illustrate correct and incorrect inference visually by treating a grayscale image of the Mona Lisa as an activity center intensity or density surface and simulating spatial capture-recapture survey data from it. Inferences can be drawn about the intensity of the point process generating activity centers, and about the likely locations of activity centers associated with the capture histories obtained from a single survey of a single realization of this process. We show that treating probabilistic predictions of activity center locations as estimates of the intensity of the process results in invalid and misleading ecological inferences, and that predictions are highly dependent on where the detectors are placed and how much survey effort is used. Estimates of the activity center density surface should be obtained by estimating the intensity of a point process model for activity centers. Practitioners should state explicitly whether they are estimating the intensity or making predictions of activity center location, and predictions of activity center locations should not be confused with estimates of the intensity.

Towards automated animal density estimation with acoustic spatial capture-recapture.

Passive acoustic monitoring can be an effective way of monitoring wildlife populations that are acoustically active but difficult to survey visually, but identifying target species calls in recordings is non-trivial. Machine learning (ML) techniques can do detection quickly but may miss calls and produce false positives, i.e., misidentify calls from other sources as being from the target species. While abundance estimation methods can address the former issue effectively, methods to deal with false positives are under-investigated. We propose an acoustic spatial capture-recapture (ASCR) method that deals with false positives by treating species identity as a latent variable. Individual-level outputs from ML techniques are treated as random variables whose distributions depend on the latent identity. This gives rise to a mixture model likelihood that we maximize to estimate call density. We compare our method to existing methods by applying it to an ASCR survey of frogs and simulated acoustic surveys of gibbons based on real gibbon acoustic data. Estimates from our method are closer to ASCR applied to the dataset without false positives than those from a widely used false positive "correction factor" method. Simulations show our method to have bias close to zero and accurate coverage probabilities and to perform substantially better than ASCR without accounting for false positives.

Counting the Capital's cats: Estimating drivers of abundance of free-roaming cats with a novel hierarchical model.

Free-roaming cats are a conservation concern in many areas but identifying their impacts and developing mitigation strategies requires a robust understanding of their distribution and density patterns. Urban and residential areas may be especially relevant in this process because free-roaming cats are abundant in these anthropogenic landscapes. Here, we estimate the occupancy and density of free-roaming cats in Washington D.C. and relate these metrics to known landscape and social factors. We conducted an extended camera trap survey of public and private spaces across D.C. and analyzed data collected from 1483 camera deployments from 2018 to 2020. We estimated citywide cat distribution by fitting hierarchical occupancy models and further estimated cat abundance using a novel random thinning spatial capture-recapture model that allows for the use of photos that can and cannot be identified to individual. Within this model, we utilized individual covariates that provided identity exclusions between photos of unidentifiable cats with inconsistent coat patterns, thus increasing the precision of abundance estimates. This combined model also allowed for unbiased estimation of density when animals cannot be identified to individual at the same rate as for free-roaming cats whose identifiability depended on their coat characteristics. Cat occupancy and abundance declined with increasing distance from residential areas, an effect that was more pronounced in wealthier neighborhoods. There was noteworthy absence of cats detected in larger public spaces and forests. Realized densities ranged from 0.02 to 1.75 cats/ha in sampled areas, resulting in a district-wide estimate of ~7296 free-roaming cats. Ninety percent of cat detections lacked collars and nearly 35% of known individuals were ear-tipped, indicative of district Trap-Neuter-Return (TNR) programs. These results suggest that we mainly sampled and estimated the unowned cat subpopulation, such that indoor/outdoor housecats were not well represented. The precise estimation of cat population densities is difficult due to the varied behavior of subpopulations within free-roaming cat populations (housecats, stray and feral cats), but our methods provide a first step in establishing citywide baselines to inform data-driven management plans for free-roaming cats in urban environments.

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.

A multidisciplinary approach to estimating wolf population size for long-term conservation.

The wolf (Canis lupus) is among the most controversial of wildlife species. Abundance estimates are required to inform public debate and policy decisions, but obtaining them at biologically relevant scales is challenging. We developed a system for comprehensive population estimation across the Italian alpine region (100,000 km2 ), involving 1513 trained operators representing 160 institutions. This extensive network allowed for coordinated genetic sample collection and landscape-level spatial capture-recapture analyses that transcended administrative boundaries to produce the first estimates of key parameters for wolf population status assessment. Wolf abundance was estimated at 952 individuals (95% credible interval 816-1120) and 135 reproductive units (i.e., packs) (95% credible interval 112-165). We also estimated that mature individuals accounted for 33-45% of the entire population. The monitoring effort was spatially estimated thereby overcoming an important limitation of citizen science data. This is an important approach for promoting wolf-human coexistence based on wolf abundance monitoring and an endorsement of large-scale harmonized conservation practices.

Una estrategia multidisciplinaria para la estimación del tamaño poblacional de los lobos para la conservación a largo plazo Resumen El lobo (Canis lupus) está entre las especies de fauna más controversiales. Se requieren estimaciones de abundancia para informar al debate público y las decisiones políticas, pero es un reto obtenerlos en escalas con relevancia biológica. Desarrollamos un sistema para la estimación completa de la población en la región alpina de Italia (100,000 km2 ), con la participación de 1,513 operadores entrenados que representan a 160 instituciones. Esta red extensa permitió una colecta coordinada de muestras genéticas y análisis de captura-recaptura espacial que trascendieron las fronteras administrativas para así producir las primeras estimaciones de los parámetros clave para la evaluación del estado de la población de los lobos. Se estimó la abundancia en 952 individuos (95% intervalo de confianza 816-1120) y 135 unidades reproductivas (es decir, manadas) (95% intervalo de confianza 112-165). También estimamos que los individuos maduros representaban el 33-45% de toda la población. El esfuerzo de monitoreo se estimó espacialmente, por lo que sobrepasó una limitación importante de la ciencia ciudadana. Esta estrategia es importante para promover la coexistencia entre lobos y humanos con base en el monitoreo de la abundancia y el apoyo a las prácticas armonizadas de conservación a gran escala.

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.

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.

Troubled spots: Human impacts constrain the density of an apex predator inside protected areas.

Effective conservation requires understanding the processes that determine population outcomes. Too often, we assume that protected areas conserve wild populations despite evidence that they frequently fail to do so. Without large-scale studies, however, we cannot determine what relationships are the product of localized conditions versus general patterns that inform conservation more broadly. Leopards' (Panthera pardus) basic ecology is well studied but little research has investigated anthropogenic effects on leopard density at broad scales. We investigated the drivers of leopard density among 27 diverse protected areas in northeastern South Africa to understand what conditions facilitate abundant populations. We formulated 10 working hypotheses that considered the relative influence of bottom-up biological factors and top-down anthropogenic factors on leopard density. Using camera-trap survey data, we fit a multi-session spatial capture-recapture model with inhomogenous density for each hypothesis and evaluated support using an information theoretic approach. The four supported hypotheses indicated that leopard density is primarily limited by human impacts, but that habitat suitability and management conditions also matter. The proportion of camera stations that recorded domestic animals, a proxy for the extent of human impacts and protected area effectiveness, was the only predictor variable present in all four supported models. Protected areas are the cornerstone of large felid conservation, but only when the human-wildlife interface is well managed and protected areas shelter wildlife populations from anthropogenic impacts. To ensure the long-term abundance of large carnivore populations, reserve managers should recognize the ineffectiveness of "paper parks" and promote contiguous networks of protected areas that offer leopards and other large mammal populations greater space and reduced human impacts.

Precision and bias of spatial capture-recapture estimates: A multi-site, multi-year Utah black bear case study.

Spatial capture-recapture (SCR) models are powerful analytical tools that have become the standard for estimating abundance and density of wild animal populations. When sampling populations to implement SCR, the number of unique individuals detected, total recaptures, and unique spatial relocations can be highly variable. These sample sizes influence the precision and accuracy of model parameter estimates. Testing the performance of SCR models with sparse empirical data sets typical of low-density, wide-ranging species can inform the threshold at which a more integrated modeling approach with additional data sources or additional years of monitoring may be required to achieve reliable, precise parameter estimates. Using a multi-site, multi-year Utah black bear (Ursus americanus) capture-recapture data set, we evaluated factors influencing the uncertainty of SCR structural parameter estimates, specifically density, detection, and the spatial scale parameter, sigma. We also provided some of the first SCR density estimates for Utah black bear populations, which ranged from 3.85 to 74.33 bears/100 km2 . Increasing total detections decreased the uncertainty of density estimates, whereas an increasing number of total recaptures and individuals with recaptures decreased the uncertainty of detection and sigma estimates, respectively. In most cases, multiple years of data were required for precise density estimates (<0.2 coefficient of variation [CV]). Across study areas there was an average decline in CV of 0.07 with the addition of another year of data. One sampled population with very high estimated bear density had an atypically low number of spatial recaptures relative to total recaptures, apparently inflating density estimates. A complementary simulation study used to assess estimate bias suggested that when <30% of recaptured individuals were spatially recaptured, density estimates were unreliable and ranged widely, in some cases to >3 times the simulated density. Additional research could evaluate these requirements for other density scenarios. Large numbers of individuals detected, numbers of spatial recaptures, and precision alone may not be sufficient indicators of parameter estimate reliability. We provide an evaluation of simple summary statistics of capture-recapture data sets that can provide an early signal of the need to alter sampling design or collect auxiliary data before model implementation to improve estimate precision and accuracy.

Estimating red fox density using non-invasive genetic sampling and spatial capture-recapture modelling.

Spatial capture-recapture modelling (SCR) is a powerful tool for estimating density, population size, and space use of elusive animals. Here, we applied SCR modelling to non-invasive genetic sampling (NGS) data to estimate red fox (Vulpes vulpes) densities in two areas of boreal forest in central (2016-2018) and southern Norway (2017-2018). Estimated densities were overall lower in the central study area (mean = 0.04 foxes per km2 in 2016, 0.10 in 2017, and 0.06 in 2018) compared to the southern study area (0.16 in 2017 and 0.09 in 2018). We found a positive effect of forest cover on density in the central, but not the southern study area. The absence of an effect in the southern area may reflect a paucity of evidence caused by low variation in forest cover. Estimated mean home-range size in the central study area was 45 km2 [95%CI 34-60] for females and 88 km2 [69-113] for males. Mean home-range sizes were smaller in the southern study area (26 km2 [16-42] for females and 56 km2 [35-91] for males). In both study areas, detection probability was session-dependent and affected by sampling effort. This study highlights how SCR modelling in combination with NGS can be used to efficiently monitor red fox populations, and simultaneously incorporate ecological factors and estimate their effects on population density and space use.

Quantifying 25 years of disease-caused declines in Tasmanian devil populations: host density drives spatial pathogen spread.

Infectious diseases are strong drivers of wildlife population dynamics, however, empirical analyses from the early stages of pathogen emergence are rare. Tasmanian devil facial tumour disease (DFTD), discovered in 1996, provides the opportunity to study an epizootic from its inception. We use a pattern-oriented diffusion simulation to model the spatial spread of DFTD across the species' range and quantify population effects by jointly modelling multiple streams of data spanning 35 years. We estimate the wild devil population peaked at 53 000 in 1996, less than half of previous estimates. DFTD spread rapidly through high-density areas, with spread velocity slowing in areas of low host densities. By 2020, DFTD occupied >90% of the species' range, causing 82% declines in local densities and reducing the total population to 16 900. Encouragingly, our model forecasts the population decline should level-off within the next decade, supporting conservation management focused on facilitating evolution of resistance and tolerance.

Experimental evaluation of spatial capture-recapture study design.

A principal challenge impeding strong inference in analyses of wild populations is the lack of robust and long-term data sets. Recent advancements in analytical tools used in wildlife science may increase our ability to integrate smaller data sets and enhance the statistical power of population estimates. One such advancement, the development of spatial capture-recapture (SCR) methods, explicitly accounts for differences in spatial study designs, making it possible to equate multiple study designs in one analysis. SCR has been shown to be robust to variation in design as long as minimal sampling guidance is adhered to. However, these expectations are based on simulation and have yet to be evaluated in wild populations. Here we conduct a rigorously designed field experiment by manipulating the arrangement of artificial cover objects (ACOs) used to collect data on red-backed salamanders (Plethodon cinereus) to empirically evaluate the effects of design configuration on inference made using SCR. Our results suggest that, using SCR, estimates of space use and detectability are sensitive to study design configuration, namely the spacing and extent of the array, and that caution is warranted when assigning biological interpretation to these parameters. However, estimates of population density remain robust to design except when the configuration of detectors grossly violates existing recommendations.

Evaluating and integrating spatial capture-recapture models with data of variable individual identifiability.

Spatial capture-recapture (SCR) models have become the preferred tool for estimating densities of carnivores. Within this family of models are variants requiring identification of all individuals in each encounter (SCR), a subset of individuals only (generalized spatial mark-resight, gSMR), or no individual identification (spatial count or spatial presence-absence). Although each technique has been shown through simulation to yield unbiased results, the consistency and relative precision of estimates across methods in real-world settings are seldom considered. We tested a suite of models ranging from those only requiring detections of unmarked individuals to others that integrate remote camera, physical capture, genetic, and global positioning system (GPS) data into a hybrid model, to estimate population densities of black bears, bobcats, cougars, and coyotes. For each species, we genotyped fecal DNA collected with detection dogs during a 20-d period. A subset of individuals from each species was affixed with GPS collars bearing unique markings and resighted by remote cameras over 140 d contemporaneous with scat collection. Camera-based gSMR models produced density estimates that differed by <10% from genetic SCR for bears, cougars, and coyotes once important sources of variation (sex or behavioral status) were controlled for. For bobcats, SCR estimates were 33% higher than gSMR. The cause of the discrepancies in estimates was likely attributable to challenges designing a study compatible for species with disparate home range sizes and the difficulty of collecting sufficient data in a timeframe in which demographic closure could be assumed. Unmarked models estimated densities that varied greatly from SCR, but estimates became more consistent in models wherein more individuals were identifiable. Hybrid models containing all data sources exhibited the most precise estimates for all species. For studies in which only sparse data can be obtained and the strictest model assumptions are unlikely to be met, we suggest researchers use caution making inference from models lacking individual identity. For best results, we further recommend the use of methods requiring at least a subset of the population is marked and that multiple data sets are incorporated when possible.

Evaluation of camera trap-based abundance estimators for unmarked populations.

Estimates of species abundance are critical to understand population processes and to assess and select management actions. However, capturing and marking individuals for abundance estimation, while providing robust information, can be economically and logistically prohibitive, particularly for species with cryptic behavior. Camera traps can be used to collect data at temporal and spatial scales necessary for estimating abundance, but the use of camera traps comes with limitations when target species are not uniquely identifiable (i.e., "unmarked"). Abundance estimation is particularly useful in the management of invasive species, with herpetofauna being recognized as some of the most pervasive and detrimental invasive vertebrate species. However, the use of camera traps for these taxa presents additional challenges with relevancy across multiple taxa. It is often necessary to use lures to attract animals in order to obtain sufficient observations, yet lure attraction can influence species' landscape use and potentially induce bias in abundance estimators. We investigated these challenges and assessed the feasibility of obtaining reliable abundance estimates using camera-trapping data on a population of invasive brown treesnakes (Boiga irregularis) in Guam. Data were collected using camera traps in an enclosed area where snakes were subject to high-intensity capture-recapture effort, resulting in presumed abundance of 116 snakes (density = 23/ha). We then applied spatial count, random encounter and staying time, space to event, and instantaneous sampling estimators to photo-capture data to estimate abundance and compared estimates to our presumed abundance. We found that all estimators for unmarked populations performed poorly, with inaccurate or imprecise abundance estimates that limit their usefulness for management in this system. We further investigated the sensitivity of these estimators to the use of lures (i.e., violating the assumption that animal behavior is unchanged by sampling) and camera density in a simulation study. Increasing the effective distances of a lure (i.e., lure attraction) and camera density both resulted in biased abundance estimates. Each estimator rarely recovered truth or suffered from convergence issues. Our results indicate that, when limited to unmarked estimators and the use of lures, camera traps alone are unlikely to produce abundance estimates with utility for brown treesnake management.

Open population maximum likelihood spatial capture-recapture.

Open population capture-recapture models are widely used to estimate population demographics and abundance over time. Bayesian methods exist to incorporate open population modeling with spatial capture-recapture (SCR), allowing for estimation of the effective area sampled and population density. Here, open population SCR is formulated as a hidden Markov model (HMM), allowing inference by maximum likelihood for both Cormack-Jolly-Seber and Jolly-Seber models, with and without activity center movement. The method is applied to a 12-year survey of male jaguars (Panthera onca) in the Cockscomb Basin Wildlife Sanctuary, Belize, to estimate survival probability and population abundance over time. For this application, inference is shown to be biased when assuming activity centers are fixed over time, while including a model for activity center movement provides negligible bias and nominal confidence interval coverage, as demonstrated by a simulation study. The HMM approach is compared with Bayesian data augmentation and closed population models for this application. The method is substantially more computationally efficient than the Bayesian approach and provides a lower root-mean-square error in predicting population density compared to closed population models.

Cluster capture-recapture to account for identification uncertainty on aerial surveys of animal populations.

Capture-recapture methods for estimating wildlife population sizes almost always require their users to identify every detected animal. Many modern-day wildlife surveys detect animals without physical capture-visual detection by cameras is one such example. However, for every pair of detections, the surveyor faces a decision that is often fraught with uncertainty: are they linked to the same individual? An inability to resolve every such decision to a high degree of certainty prevents the use of standard capture-recapture methods, impeding the estimation of animal density. Here, we develop an estimator for aerial surveys, on which two planes or unmanned vehicles (drones) fly a transect over the survey region, detecting individuals via high-definition cameras. Identities remain unknown, so one cannot discern if two detections match to the same animal; however, detections in close proximity are more likely to match. By modeling detection locations as a clustered point process, we extend recently developed methodology and propose a precise and computationally efficient estimator of animal density that does not require individual identification. We illustrate the method with an aerial survey of porpoise, on which cameras detect individuals at the surface of the sea, and we need to take account of the fact that they are not always at the surface.

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.

The enemy of my enemy is my friend: native pine marten recovery reverses the decline of the red squirrel by suppressing grey squirrel populations.

Shared enemies may instigate or modify competitive interactions between species. The dis-equilibrium caused by non-native species introductions has revealed that the outcome of such indirect interactions can often be dramatic. However, studies of enemy-mediated competition mostly consider the impact of a single enemy, despite species being embedded in complex networks of interactions. Here, we demonstrate that native red and invasive grey squirrels in Britain, two terrestrial species linked by resource and disease-mediated apparent competition, are also now linked by a second enemy-mediated relationship involving a shared native predator recovering from historical persecution, the European pine marten. Through combining spatial capture-recapture techniques to estimate pine marten density, and squirrel site-occupancy data, we find that the impact of exposure to predation is highly asymmetrical, with non-native grey squirrel occupancy strongly negatively affected by exposure to pine martens. By contrast, exposure to pine marten predation has an indirect positive effect on red squirrel populations. Pine marten predation thus reverses the well-documented outcome of resource and apparent competition between red and grey squirrels.

Dynamic occupancy modelling reveals a hierarchy of competition among fishers, grey foxes and ringtails.

Determining how species coexist is critical for understanding functional diversity, niche partitioning and interspecific interactions. Identifying the direct and indirect interactions among sympatric carnivores that enable their coexistence is particularly important to elucidate because they are integral for maintaining ecosystem function. We studied the effects of removing nine fishers (Pekania pennanti) on their population dynamics and used this perturbation to elucidate the interspecific interactions among fishers, grey foxes (Urocyon cinereoargenteus) and ringtails (Bassariscus astutus). Grey foxes (family: Canidae) are likely to compete with fishers due to their similar body sizes and dietary overlap, and ringtails (family: Procyonidae), like fishers, are semi-arboreal species of conservation concern. We used spatial capture-recapture to investigate fisher population numbers and dynamic occupancy models that incorporated interspecific interactions to investigate the effects members of these species had on the colonization and persistence of each other's site occupancy. The fisher population showed no change in density for up to 3 years following the removals of fishers for translocations. In contrast, fisher site occupancy decreased in the years immediately following the translocations. During this same time period, site occupancy by grey foxes increased and remained elevated through the end of the study. We found a complicated hierarchy among fishers, foxes and ringtails. Fishers affected grey fox site persistence negatively but had a positive effect on their colonization. Foxes had a positive effect on ringtail site colonization. Thus, fishers were the dominant small carnivore where present and negatively affected foxes directly and ringtails indirectly. Coexistence among the small carnivores we studied appears to reflect dynamic spatial partitioning. Conservation and management efforts should investigate how intraguild interactions may influence the recolonization of carnivores to previously occupied landscapes.

Pairing field methods to improve inference in wildlife surveys while accommodating detection covariance.

It is common to use multiple field sampling methods when implementing wildlife surveys to compare method efficacy or cost efficiency, integrate distinct pieces of information provided by separate methods, or evaluate method-specific biases and misclassification error. Existing models that combine information from multiple field methods or sampling devices permit rigorous comparison of method-specific detection parameters, enable estimation of additional parameters such as false-positive detection probability, and improve occurrence or abundance estimates, but with the assumption that the separate sampling methods produce detections independently of one another. This assumption is tenuous if methods are paired or deployed in close proximity simultaneously, a common practice that reduces the additional effort required to implement multiple methods and reduces the risk that differences between method-specific detection parameters are confounded by other environmental factors. We develop occupancy and spatial capture-recapture models that permit covariance between the detections produced by different methods, use simulation to compare estimator performance of the new models to models assuming independence, and provide an empirical application based on American marten (Martes americana) surveys using paired remote cameras, hair catches, and snow tracking. Simulation results indicate existing models that assume that methods independently detect organisms produce biased parameter estimates and substantially understate estimate uncertainty when this assumption is violated, while our reformulated models are robust to either methodological independence or covariance. Empirical results suggested that remote cameras and snow tracking had comparable probability of detecting present martens, but that snow tracking also produced false-positive marten detections that could potentially substantially bias distribution estimates if not corrected for. Remote cameras detected marten individuals more readily than passive hair catches. Inability to photographically distinguish individual sex did not appear to induce negative bias in camera density estimates; instead, hair catches appeared to produce detection competition between individuals that may have been a source of negative bias. Our model reformulations broaden the range of circumstances in which analyses incorporating multiple sources of information can be robustly used, and our empirical results demonstrate that using multiple field-methods can enhance inferences regarding ecological parameters of interest and improve understanding of how reliably survey methods sample these parameters.