Scaling range sizes to threats for robust predictions of risks to biodiversity

Assessments of risk to biodiversity often rely on spatial distributions of species and ecosystems. Range‐size metrics used extensively in these assessments, such as area of occupancy (AOO), are sensitive to measurement scale, prompting proposals to measure them at finer scales or at different scales based on the shape of the distribution or ecological characteristics of the biota. Despite its dominant role in red‐list assessments for decades, appropriate spatial scales of AOO for predicting risks of species’ extinction or ecosystem collapse remain untested and contentious. There are no quantitative evaluations of the scale‐sensitivity of AOO as a predictor of risks, the relationship between optimal AOO scale and threat scale, or the effect of grid uncertainty. We used stochastic simulation models to explore risks to ecosystems and species with clustered, dispersed, and linear distribution patterns subject to regimes of threat events with different frequency and spatial extent. Area of occupancy was an accurate predictor of risk (0.81<|r|<0.98) and performed optimally when measured with grid cells 0.1–1.0 times the largest plausible area threatened by an event. Contrary to previous assertions, estimates of AOO at these relatively coarse scales were better predictors of risk than finer‐scale estimates of AOO (e.g., when measurement cells are <1% of the area of the largest threat). The optimal scale depended on the spatial scales of threats more than the shape or size of biotic distributions. Although we found appreciable potential for grid‐measurement errors, current IUCN guidelines for estimating AOO neutralize geometric uncertainty and incorporate effective scaling procedures for assessing risks posed by landscape‐scale threats to species and ecosystems.     

Long-term drivers of persistence and colonization dynamics in spatially structured amphibian populations

Many organisms live in networks of local populations connected by dispersing individuals, called spatially structured populations (SSPs), where the long-term persistence of the entire network is determined by the balance between 2 processes acting at the scale of local populations: extinction and colonization. When multiple threats act on an SSP, a comparison of the different factors determining local extinctions and colonizations is essential to plan sound conservation actions. We assessed the drivers of long-term population dynamics of multiple amphibian species at the regional scale. We used dynamic occupancy models within a Bayesian framework to identify the factors determining persistence and colonization of local populations. Because connectivity among patches is fundamental to SSPs dynamics, we considered 2 measures of connectivity acting on each focal patch: incidence of the focal species and incidence of invasive crayfish. We used meta-analysis to summarize the effect of different drivers at the community level. Persistence and colonization of local populations were jointly determined by factors acting at different scales. Persistence probability was positively related to the area and the permanence of wetlands, whereas it was negatively related to occurrence of fish. Colonization probability was highest in semipermanent wetlands and in sites with a high incidence of the focal species in nearby sites, whereas it showed a negative relationship with the incidence of invasive crayfish in the landscape. By analyzing long-term data on amphibian population dynamics, we found a strong effect of some classic features commonly used in SSP studies, such as patch area and focal species incidence. The presence of an invasive non-native species at the landscape scale emerged as one of the strongest drivers of colonization dynamics, suggesting that studies on SSPs should consider different connectivity measures more frequently, such as the incidence of predators, especially when dealing with biological invasions.     

Trait synergisms and the rarity, extirpation, and extinction risk of desert fishes

Understanding the causes and consequences of species extinctions is a central goal in ecology. Faced with the difficult task of identifying those species with the greatest need for conservation, ecologists have turned to using predictive suites of ecological and life-history traits to provide reasonable estimates of species extinction risk. Previous studies have linked individual traits to extinction risk, yet the nonadditive contribution of multiple traits to the entire extinction process, from species rarity to local extirpation to global extinction, has not been examined. This study asks whether trait synergisms predispose native fishes of the Lower Colorado River Basin (USA) to risk of extinction through their effects on rarity and local extirpation and their vulnerability to different sources of threat. Fish species with "slow" life histories (e.g., large body size, long life, and delayed maturity), minimal parental care to offspring, and specialized feeding behaviors are associated with smaller geographic distribution, greater frequency of local extirpation, and higher perceived extinction risk than that expected by simple additive effects of traits in combination. This supports the notion that trait synergisms increase the susceptibility of native fishes to multiple stages of the extinction process, thus making them prone to the multiple jeopardies resulting from a combination of fewer individuals, narrow environmental tolerances, and long recovery times following environmental change. Given that particular traits, some acting in concert, may differentially predispose native fishes to rarity, extirpation, and extinction, we suggest that management efforts in the Lower Colorado River Basin should be congruent with the life-history requirements of multiple species over large spatial and temporal scales.     

Relations between Species Rarity,Vulnerability, and Range Contraction for a Beetle Group in a Densely Populated Region in the Mediterranean Biodiversity Hotspot

Rarity is often considered an indication of species extinction risk, and it is frequently used to obtain measures of species vulnerability. However, there is no strong evidence of a correlation between species vulnerability and threat. Moreover, there is no consensus about how rarity should be measured. I used a multidimensional characterization of species rarity to calculate a vulnerability index for tenebrionid beetles inhabiting an Italian region in the Mediterranean biodiversity hotspot. I used different metrics to examine 3 dimensions of rarity: species range, ecology, and population. Species with rarity values below the median were scored as rare for each dimension. I combined rarity scores into a vulnerability index. I then correlated species vulnerability with range trends (expanded vs. contracted). Different measures of the same rarity dimension were strongly correlated and produced similar vulnerability scores. This result indicates rarity‐based vulnerability estimates are slightly affected by the way a certain rarity dimension is measured. Vulnerability was correlated with range trends; species with the highest vulnerability had the strongest range contraction. However, a large number of common species also underwent range contraction in the last 50 years, and there was no clear relation between range contraction and their ecology. This indicates that in general human‐induced environmental changes affected species irrespective of their assumed vulnerability and that focusing only on rare species may severely bias perceptions of the extent of species decline. Relaciones entre Rareza de Especies, Vulnerabilidad y Contracción de Distribución Geográfica para un Grupo de Escarabajos en una Región Densamente Poblada en el Hotspot de Biodiversidad del Mediterráneo     

Use of stochastic patch occupancy models in the California red-legged frog for Bayesian inference regarding past events and future persistence

Assessing causes of population decline is critically important to management of threatened species. Stochastic patch occupancy models (SPOMs) are popular tools for examining spatial and temporal dynamics of populations when presence–absence data in multiple habitat patches are available. We developed a Bayesian Markov chain method that extends existing SPOMs by focusing on past environmental changes that may have altered occupancy patterns prior to the beginning of data collection. Using occupancy data from 3 creeks, we applied the method to assess 2 hypothesized causes of population decline—in situ die-off and residual impact of past source population loss—in the California red-legged frog. Despite having no data for the 20–30 years between the hypothetical event leading to population decline and the first data collected, we were able to discriminate among hypotheses, finding evidence that in situ die-off increased in 2 of the creeks. Although the creeks had comparable numbers of occupied segments, owing to different extinction–colonization dynamics, our model predicted an 8-fold difference in persistence probabilities of their populations to 2030. Adding a source population led to a greater predicted persistence probability than did decreasing the in situ die-off, emphasizing that reversing the deleterious impacts of a disturbance may not be the most efficient management strategy. We expect our method will be useful for studying dynamics and evaluating management strategies of many species.     

Thresholds in Songbird Occurrence in Relation to Landscape Structure

Abstract:  Theory predicts the occurrence of threshold levels of habitat in landscapes, below which ecological processes change abruptly. Simulation models indicate that below critical thresholds, fragmentation of habitat influences patch occupancy by decreasing colonization rates and increasing rates of local extinction. Uncovering such putative relationships is important for understanding the demography of species and in developing sound conservation strategies. Using segmented logistic regression, we tested for thresholds in occurrence of 15 bird species as a function of the amount of suitable habitat at multiple scales (150–2000-m radii). Suitable habitat was defined quantitatively based on previously derived, spatially explicit distribution models for each species. The occurrence of 10 out of 15 species was influenced by the amount of habitat at a landscape scale (≥500-m radius). Of these species all but one were best predicted by threshold models. Six out of nine species exhibited asymptotic thresholds; the effects of habitat loss intensified at low amounts of habitat in a landscape. Landscape thresholds ranged from 8.6% habitat to 28.7% (     = 18.5 ± 2.6%[95% CI]). For two species landscape thresholds coincided with sensitivity to fragmentation; both species were more likely to occur in large patches, but only when the amount of habitat in a landscape was low. This supports the fragmentation threshold hypothesis. Nevertheless, the occurrence of most species appeared to be unaffected by fragmentation, regardless of the amount of habitat present at landscape extents. The thresholds we identified may be useful to managers in establishing conservation targets. Our results indicate that findings of landscape-scale studies conducted in regions with relatively high proportions of habitat and low fragmentation may not be applicable in regions with low habitat proportions and high fragmentation.     

Downscaling species occupancy from coarse spatial scales

The measurement and prediction of species' populations at different spatial scales is crucial to spatial ecology as well as conservation biology. An efficient yet challenging goal to achieve such population estimates consists of recording empirical species' presence and absence at a specific regional scale and then trying to predict occupancies at finer scales. So far the majority of the methods have been based on particular species' distributional features deemed to be crucial for downscaling occupancy. However, only a minority of them have dealt explicitly with specific spatial features. Here we employ a wide class of spatial point processes, the shot noise Cox processes (SNCP), to model species occupancies at different spatial scales and show that species' spatial aggregation is crucial for predicting population estimates at fine scales starting from coarser ones. These models are formulated in continuous space and locate points regardless of the arbitrary resolution that one employs to study the spatial pattern. We compare the performances of nine models, calibrated at regional scales and demonstrate that a very simple class of SNCP, the Thomas process, is able to outperform other published models in predicting occupancies down to areas four orders of magnitude smaller than the ones employed for the parameterization. We conclude by explaining the ability of the approach to infer spatially explicit information from spatially implicit measures, the potential of the framework to combine niche and spatial models, and the possibility of reversing the method to allow upscaling.     

Spatial Scale and Determination of Species Status of the Green Frog

Although the importance of spatial scale in determining species status (abundance and distribution) is widely recognized, most ecological investigations have been conducted at local scales. Our goal was to investigate the importance of spatial scale in assessing the status of the green frog ( Rana clamitans melanota) in the center of its range in eastern North America. Using repeated surveys at 160 ponds from 1992 to 1994, we investigated patterns of occupancy, abundance, and turnover at local, sub-regional, regional, and geographic scales to determine the status of the green frog in southwestern Ontario, Canada. Patterns of pond occupancy were stable at the geographic scale. Occupancy was stable in two regions and decreased in one. At the sub-regional scale, occupancy was stable in six sub-regions, increased in one, and decreased in two. Patterns of adult abundance were stable at the geographic scale. At the regional scale trends in adult abundance were increasing, decreasing, or stable in each of three regions. At the sub-regional scale abundance was stable in three, increasing in one, decreasing in two, and no trend occurred in three sub-regions. At the local scale abundance was stable at 20% of ponds, increased at 17.8%, decreased at 14.4%, and no trend existed at 47.8%. Colonization and extinction rates ranged from 0 to 0.20 and 0 to 0.35 ponds/pond occupied/year, respectively, and differed among regions. Local extinctions took place at 25% of ponds during the study, but no sub-regional or regional scale extinctions occurred. Small populations (<10 adults/pond) were prone to local extinction. Determination of the status of the green frog is scale dependent. Although green frog populations are dynamic, it is common and stable at the geographic scale, but its status varies among regions or sub-regions. Although processes that negatively affect a species may operate at the local scale, a large-scale perspective is necessary to determine status.     

Influence of habitat quality, population size, patch size, and connectivity on patch-occupancy dynamics of the middle spotted woodpecker

Despite extensive research on the effects of habitat fragmentation, the ecological mechanisms underlying colonization and extinction processes are poorly known, but knowledge of these mechanisms is essential to understanding the distribution and persistence of populations in fragmented habitats. We examined these mechanisms through multiseason occupancy models that elucidated patch-occupancy dynamics of Middle Spotted Woodpeckers (Dendrocopos medius) in northwestern Spain. The number of occupied patches was relatively stable from 2000 to 2010 (15-24% of 101 patches occupied every year) because extinction was balanced by recolonization. Larger and higher quality patches (i.e., higher density of oaks >37 cm dbh [diameter at breast height]) were more likely to be occupied. Habitat quality (i.e., density of large oaks) explained more variation in patch colonization and extinction than did patch size and connectivity, which were both weakly associated with probabilities of turnover. Patches of higher quality were more likely to be colonized than patches of lower quality. Populations in high-quality patches were less likely to become extinct. In addition, extinction in a patch was strongly associated with local population size but not with patch size, which means the latter may not be a good surrogate of population size in assessments of extinction probability. Our results suggest that habitat quality may be a primary driver of patch-occupancy dynamics and may increase the accuracy of models of population survival. We encourage comparisons of competing models that assess occupancy, colonization, and extinction probabilities in a single analytical framework (e.g., dynamic occupancy models) so as to shed light on the association of habitat quality and patch geometry with colonization and extinction processes in different settings and species.     

Reliability of a Higher-Taxon Approach to Richness, Rarity, and Composition Assessments at the Local Scale

Abstract:  A promising shortcut for quantifying species patterns is to use genera and families as surrogates of species. At large spatial scales, concurrence between patterns of richness, rarity, and composition of species and higher taxa is generally high. Only a few researchers, however, have examined this relationship at the local scale, which is frequently the relevant scale in land-use conflicts. We investigated the reliability of the higher-taxon approach in assessing patterns of species richness, rarity, and composition at the local scale. We studied diversity patterns of three commonly used surrogate taxa: vascular plants, ground-dwelling beetles, and moths. We conducted year-round field surveys for these taxa in the Jerusalem Mountains and the Judean foothills, Israel. Richness and composition of species were highly correlated with richness and composition of genera for all taxa. At the family level, correlations with richness and composition of species were much lower. Excluding monotypic genera and families did not affect these relations. Rarity representation based on higher taxa varied considerably depending on the taxon, and rarity scale and was weaker compared with richness and composition representation. Cumulative richness curves of species and genera showed similar patterns, leveling off at equivalent sampling efforts. Genus-level assessments were a reliable surrogate for local patterns of species richness, rarity, and composition, but family-level assessments performed poorly. The advantage of using coarse taxonomic scales in local diversity surveys is that it may decrease identification time and the need for experts, but it will not reduce sampling effort.     

Transient dynamics of invasive competition: barred owls, spotted owls, habitat, and the demons of competition present

The recent range expansion of Barred Owls (Strix varia) into the Pacific Northwest, where the species now co-occurs with the endemic Northern Spotted Owl (Strix occidentalis caurina), resulted in a unique opportunity to investigate potential competition between two congeneric, previously allopatric species. The primary criticism of early competition research was the use of current species' distribution patterns to infer past processes; however, the recent expansion of the Barred Owl and the ability to model the processes that result in site occupancy (i.e., colonization and extinction) allowed us to address the competitive process directly rather than inferring past processes through current patterns. The purpose of our study was to determine whether Barred Owls had any negative effects on occupancy dynamics of nesting territories by Northern Spotted Owls and how these effects were influenced by habitat characteristics of Spotted Owl territories. We used single-species, multi-season occupancy models and covariates quantifying Barred Owl detections and habitat characteristics to model extinction and colonization rates of Spotted Owl pairs in southern Oregon, USA. We observed a strong, negative association between Barred Owl detections and colonization rates and a strong positive effect of Barred Owl detections on extinction rates of Spotted Owls. We observed increased extinction rates in response to decreased amounts of old forest at the territory core and higher colonization rates when old-forest habitat was less fragmented. Annual site occupancy for pairs reflected the strong effects of Barred Owls on occupancy dynamics with much lower occupancy rates predicted for territories where Barred Owls were detected. The strong Barred Owl and habitat effects on occupancy dynamics of Spotted Owls provided evidence of interference competition between the species. These effects increase the importance of conserving large amounts of contiguous, old-forest habitat to maintain Northern Spotted Owls in the landscape.     

Predicting and setting conservation priorities for Bolivian mammals based on biological correlates of the risk of decline

The recognition that growing proportions of species worldwide are endangered has led to the development of comparative analyses to elucidate why some species are more prone to extinction than others. Understanding factors and patterns of species vulnerability might provide an opportunity to develop proactive conservation strategies. Such comparative analyses are of special concern at national scales because this is the scale at which most conservation initiatives take place. We applied powerful ensemble learning models to test for biological correlates of the risk of decline among the Bolivian mammals to understand species vulnerability at a national scale and to predict the population trend for poorly known species. Risk of decline was nonrandomly distributed: higher proportions of large‐sized taxa were under decline, whereas small‐sized taxa were less vulnerable. Body mass, mode of life (i.e., aquatic, terrestrial, volant), geographic range size, litter size, home range, niche specialization, and reproductive potential were strongly associated with species vulnerability. Moreover, we found interacting and nonlinear effects of key traits on the risk of decline of mammals at a national scale. Our model predicted 35 data‐deficient species in decline on the basis of their biological vulnerability, which should receive more attention in order to prevent their decline. Our results highlight the relevance of comparative analysis at relatively narrow geographical scales, reveal previously unknown factors related to species vulnerability, and offer species‐by‐species outcomes that can be used to identify targets for conservation, especially for insufficiently known species. Predección y Definición de Prioridades de Conservación para Mamíferos de Bolivia con Base en Correlaciones Biológicas del Riesgo de Declinación     

Hierarchical analysis of species distributions and abundance across environmental gradients

Abiotic and biotic processes operate at multiple spatial and temporal scales to shape many ecological processes, including species distributions and demography. Current debate about the relative roles of niche-based and stochastic processes in shaping species distributions and community composition reflects, in part, the challenge of understanding how these processes interact across scales. Traditional statistical models that ignore autocorrelation and spatial hierarchies can result in misidentification of important ecological covariates. Here, we demonstrate the utility of a hierarchical modeling framework for testing hypotheses about the importance of abiotic factors at different spatial scales and local spatial autocorrelation for shaping species distributions and abundances. For the two orchid species studied, understory light availability and soil moisture helped to explain patterns of presence and abundance at a microsite scale (<4 m2), while soil organic content was important at a population scale (<400 m2). The inclusion of spatial autocorrelation is shown to alter the magnitude and certainty of estimated relationships between abundance and abiotic variables, and we suggest that such analysis be used more often to explore the relationships between species life histories and distributions. The hierarchical modeling framework is shown to have great potential for elucidating ecological relationships involving abiotic and biotic processes simultaneously at multiple scales.     

Testing Hypotheses of Bird Extinctions at Rio Palenque,Ecuador, with Informal Species Lists

Abstract: Informally gathered species lists are a potential source of data for conservation biology, but most remain unused because of questions of reliability and statistical issues. We applied two alternative analytical methods (contingency tests and occupancy modeling) to a 35‐year data set (1973–2007) to test hypotheses about local bird extinction. We compiled data from bird lists collected by expert amateurs and professional scientists in a 2‐km² fragment of lowland tropical forest in coastal Ecuador. We tested the effects of the following on local extinction: trophic level, sociality, foraging specialization, light tolerance, geographical range area, and biogeographic source. First we assessed extinction on the basis of the number of years in which a species was not detected on the site and used contingency tests with each factor to compare the frequency of expected and observed extinction events among different species categories. Then we defined four multiyear periods that reflected different stages of deforestation and isolation of the study site and used occupancy modeling to test extinction hypotheses singly and in combination. Both types of analyses supported the biogeographic source hypothesis and the species‐range hypothesis as causes of extinction; however, occupancy modeling indicated the model incorporating all factors except foraging specialization best fit the data.     

Quantification of Extinction Risk: IUCN's System for Classifying Threatened Species

Abstract: The International Union for Conservation of Nature (IUCN) Red List of Threatened Species was increasingly used during the 1980s to assess the conservation status of species for policy and planning purposes. This use stimulated the development of a new set of quantitative criteria for listing species in the categories of threat: critically endangered, endangered, and vulnerable. These criteria, which were intended to be applicable to all species except microorganisms, were part of a broader system for classifying threatened species and were fully implemented by IUCN in 2000. The system and the criteria have been widely used by conservation practitioners and scientists and now underpin one indicator being used to assess the Convention on Biological Diversity 2010 biodiversity target. We describe the process and the technical background to the IUCN Red List system. The criteria refer to fundamental biological processes underlying population decline and extinction. But given major differences between species, the threatening processes affecting them, and the paucity of knowledge relating to most species, the IUCN system had to be both broad and flexible to be applicable to the majority of described species. The system was designed to measure the symptoms of extinction risk, and uses 5 independent criteria relating to aspects of population loss and decline of range size. A species is assigned to a threat category if it meets the quantitative threshold for at least one criterion. The criteria and the accompanying rules and guidelines used by IUCN are intended to increase the consistency, transparency, and validity of its categorization system, but it necessitates some compromises that affect the applicability of the system and the species lists that result. In particular, choices were made over the assessment of uncertainty, poorly known species, depleted species, population decline, restricted ranges, and rarity; all of these affect the way red lists should be viewed and used. Processes related to priority setting and the development of national red lists need to take account of some assumptions in the formulation of the criteria.     

The Effect of Fragment Shape and Species' Sensitivity to Habitat Edges on Animal Population Size

Abstract:  Habitat fragmentation causes extinction of local animal populations by decreasing the amount of viable "core" habitat area and increasing edge effects. It is widely accepted that larger fragments make better nature reserves because core-dwelling species have a larger amount of suitable habitat. Nevertheless, fragments in real landscapes have complex, irregular shapes. We modeled the population sizes of species that have a representative range of preferences for or aversions to habitat edges at five spatial scales (within 10, 32, 100, 320, and 1000 m of an edge) in a nation-wide analysis of forest remnants in New Zealand. We hypothesized that the irregular shapes of fragments in real landscapes should generate statistically significant correlations between population density and fragment area, purely as a "geometric" effect of varying species responses to the distribution of edge habitat. Irregularly shaped fragments consistently reduced the population size of core-dwelling species by 10–100%, depending on the scale over which species responded to habitat edges. Moreover, core populations within individual fragments were spatially discontinuous, containing multiple, disjunct populations that inhabited small spatial areas and had reduced population size. The geometric effect was highly nonlinear and depended on the range of fragment sizes sampled and the scale at which species responded to habitat edges. Fragment shape played a strong role in determining population size in fragmented landscapes; thus, habitat restoration efforts may be more effective if they focus on connecting disjunct cores rather than isolated fragments.     

Effects of Recent Environmental Change on Accuracy of Inferences of Extinction Status

Correctly classifying a species as extinct or extant is of critical importance if current rates of biodiversity loss are to be accurately quantified. Observing an extinction event is rare, so in many cases extinction status is inferred using methods based on the analysis of records of historic sighting events. The accuracy of such methods is difficult to test. However, results of recent experiments with microcosm communities suggest that the rate at which a population declines to extinction, potentially driven by varying environmental conditions, may alter one's ability accurately to infer extinction status. We tested how the rate of population decline, driven by historic environmental change, alters the accuracy of 6 commonly applied sighting‐based methods used to infer extinction. We used data from small‐scale experimental communities and recorded wild population extirpations. We assessed how accuracy of the different methods was affected by rate of population decline, search effort, and number of sighting events recorded. Rate of population decline and historic population size of the species affected the accuracy of inferred extinction dates; however, faster declines produced more accurate inferred dates of extinction, but only when population sizes were higher. Optimal linear estimation (OLE) offered the most reliable and robust estimates, though no single method performed best in all situations, and it may be appropriate to use a different method if information regarding historic search efforts is available. OLE provided the most accurate estimates of extinction when the number of sighting events used was >10, and future use of this method should take this into account. Data from experimental populations provide added insight into testing techniques to discern wild extirpation events. Care should be taken designing such experiments so that they mirror closely the abundance dynamics of populations affected by real‐world extirpation events. Efectos del Cambio Ambiental Reciente sobre la Precisión de las Inferencias sobre el Estado de Extinción     

Area-based assessment of extinction risk

Underpinning the International Union for Conservation of Nature (IUCN) Red List is the assessment of extinction risk as determined by the size and degree of loss of populations. The IUCN system lists a species as Critically Endangered, Endangered, or Vulnerable if its population size declines 80%, 50%, or 30% within a given time frame. However, effective implementation of the system faces substantial challenges and uncertainty because geographic scale data on population size and long-term dynamics are scarce. I develop a model to quantify extinction risk using a measure based on a species' distribution, a much more readily obtained quantity. The model calculates the loss of the area of occupancy that is equivalent to the loss of a given proportion of a population. It is a very simple yet general model that has no free parameters and is independent of scale. The model predicted well the distributions of 302 tree species at a local scale and the distributions of 348 species of North American land birds. This area-based model provides a solution to the long-standing problem for IUCN assessments of lack of data on population sizes, and thus it will contribute to facilitating the quantification of extinction risk worldwide.     

A model for behavioral regulation of metapopulation dynamics

Habitat fragmentation can decrease local population persistence by reducing connectivity, which is a function of dispersal of individuals among habitat fragments. Dispersal is often treated as diffusion in population models, even though for many species it is a result of a series of behavioral decisions. We developed a metapopulation model to explore the potential importance of dispersal behaviors in driving metapopulation dynamics. We incorporated types of behavior that affect dispersal—colonization inhibiting, colonization enhancing, extinction inhibiting, extinction enhancing, rescue enhancing, rescue inhibiting—into Levins’ (1969) metapopulation model and projected occupancy rates for a variety of parameter values. Examples from the literature of behaviors associated with each of these parameters are provided. Our model simplifies into previously published metapopulation models that incorporate only a single behavior, and we present a density-dependent rescue function that leads to multiple non-zero equilibria. We found a variety of behavioral effects on metapopulations. Rescue enhancement fills patches faster than does colonization enhancement or extinction inhibition, and declines in patch occupancy are moderate with extinction enhancement, but colonization inhibition causes metapopulation extinction. We also found that with colonization and extinction inhibitions, equilibrium patch occupancy is inversely related to patch turnover rate. With density-dependent rescue, persistence depends not only on the strength of the strong rescue effect, but also on having a sufficient initial fraction of patches occupied; the stronger the rescue effect, the lower this fraction can be. This study suggests that dispersal behavior can have strong influences on metapopulation dynamics. It confirms the importance of understanding the relationship between landscape structure and dispersal behavior in understanding population persistence.     

Effect of scale on trait predictors of species responses to agriculture

Species persistence in human‐altered landscapes can depend on factors operating at multiple spatial scales. To understand anthropogenic impacts on biodiversity, it is useful to examine relationships between species traits and their responses to land‐use change. A key knowledge gap concerns whether these relationships vary depending on the scale of response under consideration. We examined how local‐ and large‐scale habitat variables influence the occupancy dynamics of a bird community in cloud forest zones in the Colombian Chocó‐Andes. Using data collected across a continuum of forest and agriculture, we examined which traits best predict species responses to local variation in farmland and which traits best predict species responses to isolation from contiguous forest. Global range size was a strong predictor of species responses to agriculture at both scales; widespread species were less likely to decline as local habitat cover decreased and as distance from forest increased. Habitat specialization was a strong predictor of species responses only at the local scale. Open‐habitat species were particularly likely to increase as pasture increased, but they were relatively insensitive to variation in distance to forest. Foraging plasticity and flocking behavior were strong predictors of species responses to distance from forest, but not their responses to local habitat. Species with lower plasticity in foraging behaviors and obligate flock‐following species were more likely to decline as distance from contiguous forest increased. For species exhibiting these latter traits, persistence in tropical landscapes may depend on the protection of larger contiguous blocks of forest, rather than the integration of smaller‐scale woodland areas within farmland. Species listed as threatened or near threatened on the International Union for Conservation of Nature Red List were also more likely to decline in response to both local habitat quality and isolation from forest relative to least‐concern species, underlining the importance of contiguous forests for threatened taxa.