Neighborhood and habitat effects on vital rates: expansion of the Barred Owl in the Oregon coast ranges

In this paper, we modify dynamic occupancy models developed for detection-nondetection data to allow for the dependence of local vital rates on neighborhood occupancy, where neighborhood is defined very flexibly. Such dependence of occupancy dynamics on the status of a relevant neighborhood is pervasive, yet frequently ignored. Our framework permits joint inference about the importance of neighborhood effects and habitat covariates in determining colonization and extinction rates. Our specific motivation is the recent expansion of the Barred Owl (Strix varia) in western Oregon, USA, over the period 1990-2010. Because the focal period was one of dramatic range expansion and local population increase, the use of models that incorporate regional occupancy (sources of colonists) as determinants of dynamic rate parameters is especially appropriate. We began our analysis of 21 years of Barred Owl presence/nondetection data in the Tyee Density Study Area (TDSA) by testing a suite of six models that varied only in the covariates included in the modeling of detection probability. We then tested whether models that used regional occupancy as a covariate for colonization and extinction outperformed models with constant or year-specific colonization or extinction rates. Finally we tested whether habitat covariates improved the AIC of our models, focusing on which habitat covariates performed best, and whether the signs of habitat effects are consistent with a priori hypotheses. We conclude that all covariates used to model detection probability lead to improved AIC, that regional occupancy influences colonization and extinction rates, and that habitat plays an important role in determining extinction and colonization rates. As occupancy increases from low levels toward equilibrium, colonization increases and extinction decreases, presumably because there are more and more dispersing juveniles. While both rates are affected, colonization increases more than extinction decreases. Colonization is higher and extinction is lower in survey polygons with more riparian forest. The effects of riparian forest on extinction rates are greater than on colonization rates. Model results have implications for management of the invading Barred Owl, both through habitat alteration and removal.     

Genetic Identification of Spotted Owls, Barred Owls, and Their Hybrids: Legal Implications of Hybrid Identity

Abstract:  Recent population expansion of Barred Owls (  Strix varia ) into western North America has led to concern that they may compete with and further harm the Northern Spotted Owl (  S. occidentalis caurina ), which is already listed as threatened under the U.S. Endangered Species Act (ESA). Because they hybridize, there is a legal need under the ESA for forensic identification of both species and their hybrids. We used mitochondrial control-region DNA and amplified fragment-length polymorphism (AFLP) analyses to assess maternal and biparental gene flow in this hybridization process. Mitochondrial DNA sequences (524 base pairs) indicated large divergence between Barred and Spotted Owls (13.9%). Further, the species formed two distinct clades with no signs of previous introgression. Fourteen diagnostic AFLP bands also indicated extensive divergence between the species, including markers differentiating them. Principal coordinate analyses and assignment tests clearly supported this differentiation. We found that hybrids had unique genetic combinations, including AFLP markers from both parental species, and identified known hybrids as well as potential hybrids with unclear taxonomic status. Our analyses corroborated the findings of extensive field studies that most hybrids genetically sampled resulted from crosses between female Barred Owls and male Spotted Owls. These genetic markers make it possible to clearly identify these species as well as hybrids and can now be used for research, conservation, and law enforcement. Several legal avenues may facilitate future conservation of Spotted Owls and other ESA-listed species that hybridize, including the ESA similarity-of-appearance clause (section 4[e]) and the Migratory Bird Treaty Act. The Migratory Bird Treaty Act appears to be the most useful route at this time.     

Conservation of the Northern Spotted Owl under the Northwest Forest Plan

Abstract:  Development of the Northwest Forest Plan (NWFP) was motivated by concerns about the overharvest of late-seral forests and the effects of intensive forest management on the long-term viability of the Northern Spotted Owl ( Strix occidentalis caurina ). Following several years of intense political and legal debates, the final NWFP was approved in 1994. Even though the plan evolved with a broad ecosystem perspective, it remained anchored in the Spotted Owl reserve design proposed in 1990. Based on a criterion of stable or increasing populations, a decade later it remains unclear whether the enactment of the NWFP has improved the conservation status of Spotted Owls. The results of intensive monitoring of several Spotted Owl populations for over a decade suggest a continuing range-wide decline even though rates of timber harvest have declined dramatically on federal lands. The cause of the decline is difficult to determine because the research needed to establish cause and effect relations has not been done. One plausible hypothesis is that the owl's life history greatly constrains its rate of population growth even when habitat is no longer limiting. Since enactment of the NWFP, new threats have arisen, including the movement of Barred Owls ( S. varia ) into the range of the Spotted Owl, political pressure to increase levels of timber harvest, and recent changes to forest laws that eliminate the requirement to assess the viability of wildlife populations on U.S. Department of Agriculture Forest Service lands. At this time is appears that Spotted Owl conservation rests critically on continued implementation of the protections afforded by the NWFP and the U.S. Endangered Species Act.     

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.     

Overestimation of Fire Risk in the Northern Spotted Owl Recovery Plan

Abstract:  The U.S. Fish and Wildlife Service's recent recovery plan for one of the most carefully watched threatened species worldwide, the Northern Spotted Owl (Strix occidentalis caurina), recommended a major departure in conservation strategies in the northwestern United States. Due to concern about fire, the plan would switch from a reserve to a no-reserve strategy in up to 52% of the owl's range. Fuel treatments (e.g., thinning) at regular intervals also would occur on up to 65–70% of dry forests in this area. Estimations of fire risk, however, were based on less than a decade of data and an anecdotal assessment of a single, large fire. We found that decadal data are inherently too short, given infrequent large fires, to accurately predict fire risk and trends. Rates of high-severity fire, based on remote-sensing data, are far lower than reported in the plan and in comparison with the rate of old-forest recruitment. In addition, over a 22-year period, there has been no increase in the proportion of high-severity fire. Our findings refute the key conclusions of the plan that are the basis for major changes in conservation strategies for the Spotted Owl. The best available science is needed to address these strategies in an adaptive-management framework. From the standpoint of fire risk, there appears to be ample time for research on fire and proposed treatment effects on Spotted Owls before designing extensive management actions or eliminating reserves.     

Evaluating Areas of High Conservation Value in Western Oregon with a Decision‐Support Model

Abstract: The Northwest Forest Plan was implemented in 1994 to protect habitat for species associated with old‐growth forests, including Northern Spotted Owls (Strix occidentailis caurina) in Washington, Oregon, and northern California (U.S.A.). Nevertheless, 10‐year monitoring data indicate mixed success in meeting the ecological goals of the plan. We used the ecosystem management decision‐support model to evaluate terrestrial and aquatic habitats across the landscape on the basis of ecological objectives of the Northwest Forest Plan, which included maintenance of late‐successional and old‐growth forest, recovery, and maintenance of Pacific salmon (Oncorhynchus spp.), and viability of Northern Spotted Owls. Areas of the landscape that contained habitat characteristics that supported these objectives were considered of high conservation value. We used the model to evaluate ecological condition of each of the 36, 180 township and range sections of the study area. Eighteen percent of the study area was identified as habitat of high conservation value. These areas were mostly on public lands. Many of the sections that contained habitat of exceptional conservation value were on Bureau of Land Management land that has been considered for management‐plan revisions to increase timber harvests. The results of our model can be used to guide future land management in the Northwest Forest Plan area, and illustrate how decision‐support models can help land managers develop strategies to better meet their goals.     

Divergence in sink contributions to population persistence

Population sinks present unique conservation challenges. The loss of individuals in sinks can compromise persistence; but conversely, sinks can improve viability by improving connectivity and facilitating the recolonization of vacant sources. To assess the contribution of sinks to regional population persistence of declining populations, we simulated source–sink dynamics for 3 very different endangered species: Black‐capped Vireos (Vireo atricapilla) at Fort Hood, Texas, Ord's kangaroo rats (Dipodomys ordii) in Alberta, and Northern Spotted Owls (Strix occidentalis caurina) in the northwestern United States. We used empirical data from these case studies to parameterize spatially explicit individual‐based models. We then used the models to quantify population abundance and persistence with and without long‐term sinks. The contributions of sink habitats varied widely. Sinks were detrimental, particularly when they functioned as strong sinks with few emigrants in declining populations (e.g., Alberta's Ord's kangaroo rat) and benign in robust populations (e.g., Black‐capped Vireos) when Brown‐headed Cowbird (Molothrus ater) parasitism was controlled. Sinks, including ecological traps, were also crucial in delaying declines when there were few sources (e.g., in Black‐capped Vireo populations with no Cowbird control). Sink contributions were also nuanced. For example, sinks that supported large, variable populations were subject to greater extinction risk (e.g., Northern Spotted Owls). In each of our case studies, new context‐dependent sinks emerged, underscoring the dynamic nature of sources and sinks and the need for frequent re‐assessment. Our results imply that management actions based on assumptions that sink habitats are generally harmful or helpful risk undermining conservation efforts for declining populations.     

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.     

Occupancy estimation and modeling with multiple states and state uncertainty

The distribution of a species over space is of central interest in ecology, but species occurrence does not provide all of the information needed to characterize either the well-being of a population or the suitability of occupied habitat. Recent methodological development has focused on drawing inferences about species occurrence in the face of imperfect detection. Here we extend those methods by characterizing occupied locations by some additional state variable (e.g., as producing young or not). Our modeling approach deals with both detection probabilities <1 and uncertainty in state classification. We then use the approach with occupancy and reproductive rate data from California Spotted Owls (Strix occidentalis occidentalis) collected in the central Sierra Nevada during the breeding season of 2004 to illustrate the utility of the modeling approach. Estimates of owl reproductive rate were larger than na?ve estimates, indicating the importance of appropriately accounting for uncertainty in detection and state classification.     

Mitigation of Habitat "Take": Application to Habitat Conservation Planning

One of the most important provisions of the U.S. Endangered Species Act precludes the "taking" of listed species on both public and private land. In past Endangered Species Act litigation, take has been broadly interpreted to include the destruction or modification of habitats as well as the direct killing of animals. This requirement created an extensive burden on private landowners to provide habitats for listed species. This burden was substantially lessened when the ESA was modified in 1982 to allow incidental takings conditioned on preparation of a satisfactory "habitat conservation plan." Because the majority of listed species are imperiled due to habitat modification, most habitat conservation plans must demonstrate defensible methods to mitigate against incidental habitat loss. A review of HCPs for the Northern Spotted Owl ( Strix occidentalis), and other species, indicates that mitigation solutions are often arbitrary, lacking an empirical foundation in the species' life history requirements. Based on data from the Spotted Owl, we illustrate a biologically based method for estimating the areal requirements necessary to mitigate against the take of essential habitats. Toward this goal we adopt the concept of "core area," that portion of an animal's home range that receives disproportionate use. We estimated core areas by means of the adaptive kernel density function and tested against a null distribution of animal use that assumes a bivariate, uniform distribution of locations within the home range. The method we illustrate, which is defensible, repeatable, and empirical, is a clear improvement over the ad hoc methods used in many habitat conservation plans. Further, the methods we propose should be applicable to a large number of terrestrial species for which home range is a meaningful concept.     

Colonization and extinction in dynamic habitats: an occupancy approach for a Great Plains stream fish assemblage

Despite the importance of habitat in determining species distribution and persistence, habitat dynamics are rarely modeled in studies of metapopulations. We used an integrated habitat-occupancy model to simultaneously quantify habitat change, site fidelity, and local colonization and extinction rates for larvae of a suite of Great Plains stream fishes in the Arikaree River, eastern Colorado, USA, across three years. Sites were located along a gradient of flow intermittency and groundwater connectivity. Hydrology varied across years: the first and third being relatively wet and the second dry. Despite hydrologic variation, our results indicated that site suitability was random from one year to the next. Occupancy probabilities were also independent of previous habitat and occupancy state for most species, indicating little site fidelity. Climate and groundwater connectivity were important drivers of local extinction and colonization, but the importance of groundwater differed between periods. Across species, site extinction probabilities were highest during the transition from wet to dry conditions (range: 0.52-0.98), and the effect of groundwater was apparent with higher extinction probabilities for sites not fed by groundwater. Colonization probabilities during this period were relatively low for both previously dry sites (range: 0.02-0.38) and previously wet sites (range: 0.02-0.43). In contrast, no sites dried or remained dry during the transition from dry to wet conditions, yielding lower but still substantial extinction probabilities (range: 0.16-0.63) and higher colonization probabilities (range: 0.06-0.86), with little difference among sites with and without groundwater. This approach of jointly modeling both habitat change and species occupancy will likely be useful to incorporate effects of dynamic habitat on metapopulation processes and to better inform appropriate conservation actions.     

Consumptive and nonconsumptive effects of predators on metacommunities of competing prey

Although predators affect prey both via consumption and by changing prey migration behavior, the interplay between these two effects is rarely incorporated into spatial models of predator-prey dynamics and competition among prey. We develop a model where generalist predators have consumptive effects (i.e., altering the likelihood of local prey extinction) as well as nonconsumptive effects (altering the likelihood of colonization) on spatially separated prey populations (metapopulations). We then extend this model to explore the effects of predators on competition among prey. We find that generalist predators can promote persistence of prey metapopulations by promoting prey colonization, but predators can also hasten system-wide extinction by either increasing local extinction or reducing prey migration. By altering rates of prey migration, predators in one location can exert remote control over prey dynamics in another location via predator-mediated changes in prey flux. Thus, the effect of predators may extend well beyond the proportion of patches they visit. In the context of prey metacommunities, predator-mediated shifts in prey migration and mortality can shift the competition-colonization trade-off among competing prey, leading to changes in the prey community as well as changes in the susceptibility of prey species to habitat loss. Consequently, native prey communities may be susceptible to invasion not only by exotic prey species that experience reduced amounts of mortality from resident predators, but also by exotic prey species that exhibit strong dispersal in response to generalist native predators. Ultimately, our work suggests that the consumptive and nonconsumptive effects of generalist predators may have strong, yet potentially cryptic, effects on competing prey capable of mediating coexistence, fostering invasion, and interacting with anthropogenic habitat alteration.     

Spotted Owls and Old Growth Logging in the Pacific Northwest

Abstract: Northern Spotted Owls, Strix occidentalis caurina, require large tracts of old-growth conifer forest to survive and reproduce. Much of this forest has been or is being cut by commercial logging operations, with uncertain consequences for the owls. In this paper I present simulation models of owl population change over the next 100 years, as summing a variety of scenarios for habitat destruction and fragmentation. My analysis differs from previous models by incorporating patchy territory distribution and random environmental fluctuations. Fragmented and patchy habitat distributions are common problems for endangered organisms, but they have received little attention from modelers. My results indicate that yearly fluctuations in breeding success have little impact on owl populations, but that spatial structure is quite important and should be considered in planning forest preservation. The simulations suggest that for all reasonable parameter values the proposed US. Forest Service logging plans will lead to the demise of the owls.     

Extinction debt of forest plants persists for more than a century following habitat fragmentation

Following habitat fragmentation individual habitat patches may lose species over time as they pay off their "extinction debt." Species with relatively low rates of population extinction and colonization ("slow" species) may maintain extinction debts for particularly prolonged periods, but few data are available to test this prediction. We analyzed two unusually detailed data sets on forest plant distributions and land-use history from Lincolnshire, United Kingdom, and Vlaams-Brabant, Belgium, to test for an extinction debt in relation to species-specific extinction and colonization rates. Logistic regression models predicting the presence-absence of 36 plant species were first parameterized using data from Lincolnshire, where forest cover has been relatively low (approximately 5-8%) for the past 1000 years. Consistent with extinction debt theory, for relatively slow species (but not fast species) these models systematically underpredicted levels of patch occupancy in Vlaams-Brabant, where forest cover was reduced from approximately 25% to <10% between 1775 and 1900 (it is presently 6.5%). As a consequence, the ability of the Lincolnshire models to predict patch occupancy in Vlaams-Brabant was worse for slow than for fast species. Thus, more than a century after forest fragmentation reached its current level an extinction debt persists for species with low rates of population turnover.     

Demography of Two Mexican Spotted Owl Populations

Abstract: The Mexican Spotted Owl ( Strix occidentalis lucida ) is a threatened subspecies of the southwestern United States and northern Mexico. Although threatened, little is known about its population status. We studied the demography of the Mexican Spotted Owl in one study area in Arizona and one in New Mexico from 1991 to 1997. We estimated annual survival rates (φ), fecundity rates (   b ), and abundance (   N ) to test the hypothesis that population trends were stationary. Although annual fecundity rates (   b ^ = 0.494 for Arizona and 0.380 for New Mexico) and annual juvenile survival rates ( φ^ = 0.179 for Arizona and 0.109 for New Mexico) differed in magnitude between the study areas, they exhibited similar temporal patterns. Annual survival for territorial owls varied randomly in Arizona but declined linearly in New Mexico. Mean annual survival for territorial owls was 0.814 in Arizona and 0.832 for owls ≥3 years old and 0.644 for owls 1–2 years old in New Mexico. Based on survival and fecundity estimates, the annual rates of change (λ^) indicated that both populations were declining at ≥10% a year. These estimates were corroborated by observed declines in abundance. Some regional factor may have been affecting fecundity, whereas a combination of factors may have been affecting survival. Two possible reasons for the population declines are declines in habitat quality and regional trends in climate.     

Value of protected areas to avian persistence across 20 years of climate and land-use change

Establishing protected areas, where human activities and land cover changes are restricted, is among the most widely used strategies for biodiversity conservation. This practice is based on the assumption that protected areas buffer species from processes that drive extinction. However, protected areas can maintain biodiversity in the face of climate change and subsequent shifts in distributions have been questioned. We evaluated the degree to which protected areas influenced colonization and extinction patterns of 97 avian species over 20 years in the northeastern United States. We fitted single-visit dynamic occupancy models to data from Breeding Bird Atlases to quantify the magnitude of the effect of drivers of local colonization and extinction (e.g., climate, land cover, and amount of protected area) in heterogeneous landscapes that varied in the amount of area under protection. Colonization and extinction probabilities improved as the amount of protected area increased, but these effects were conditional on landscape context and species characteristics. In this forest-dominated region, benefits of additional land protection were greatest when both forest cover in a grid square and amount of protected area in neighboring grid squares were low. Effects did not vary with species’ migratory habit or conservation status. Increasing the amounts of land protection benefitted the range margins species but not the core range species. The greatest improvements in colonization and extinction rates accrued for forest birds relative to open-habitat or generalist species. Overall, protected areas stemmed extinction more than they promoted colonization. Our results indicate that land protection remains a viable conservation strategy despite changing habitat and climate, as protected areas both reduce the risk of local extinction and facilitate movement into new areas. Our findings suggest conservation in the face of climate change favors creation of new protected areas over enlarging existing ones as the optimal strategy to reduce extinction and provide stepping stones for the greatest number of species.     

Four Factors Modifying the Effect of Competition on Carnivore Population Dynamics as Illustrated by African Wild Dogs

Abstract: Limitation of predator populations by prey availability and the effects of predators on prey populations are widely recognized as important ecological processes that affect carnivore conservation. Interspecific competition can also be a strong limiting factor for carnivore populations, and the effects of competition help explain why some carnivore species are prone to extinction. Competition among carnivores is unusual in some ways, so some predictions from traditional models of competition do not hold. For example, an increase in the density of prey can increase the effect of competition among carnivores, rather than weakening it. I used published data from African wild dogs (    Lycaon pictus ) to highlight four complexities that can modify the effects of competition on the population dynamics of carnivores: habitat fragmentation, counterintuitive effects of prey density, predator-prey size ratios, and habitat type.     

The Northwest Forest Plan: Origins, Components, Implementation Experience, and Suggestions for Change

Abstract:  In the 1990s the federal forests in the Pacific Northwest underwent the largest shift in management focus since their creation, from providing a sustained yield of timber to conserving biodiversity, with an emphasis on endangered species. Triggered by a legal challenge to the federal protection strategy for the Northern Spotted Owl ( Strix occidentalis caurina ), this shift was facilitated by a sequence of science assessments that culminated in the development of the Northwest Forest Plan. The plan, adopted in 1994, called for an extensive system of late-successional and riparian reserves along with some timber harvest on the intervening lands under a set of controls and safeguards. It has proven more successful in stopping actions harmful to conservation of old-growth forests and aquatic systems than in achieving restoration goals and economic and social goals. We make three suggestions that will allow the plan to achieve its goals: (1) recognize that the Northwest Forest Plan has evolved into an integrative conservation strategy, (2) conserve old-growth trees and forests wherever they occur, and (3) manage federal forests as dynamic ecosystems.     

Minimum viable metapopulation size, extinction debt, and the conservation of a declining species.

A key question facing conservation biologists is whether declines in species' distributions are keeping pace with landscape change, or whether current distributions overestimate probabilities of future persistence. We use metapopulations of the marsh fritillary butterfly Euphydryas aurinia in the United Kingdom as a model system to test for extinction debt in a declining species. We derive parameters for a metapopulation model (incidence function model, IFM) using information from a 625-km2 landscape where habitat patch occupancy, colonization, and extinction rates for E. aurinia depend on patch connectivity, area, and quality. We then show that habitat networks in six extant metapopulations in 16-km2 squares were larger, had longer modeled persistence times (using IFM), and higher metapopulation capacity (lambdaM) than six extinct metapopulations. However, there was a > 99% chance that one or more of the six extant metapopulations would go extinct in 100 years in the absence of further habitat loss. For 11 out of 12 networks, minimum areas of habitat needed for 95% persistence of metapopulation simulations after 100 years ranged from 80 to 142 ha (approximately 5-9% of land area), depending on the spatial location of habitat. The area of habitat exceeded the estimated minimum viable metapopulation size (MVM) in only two of the six extant metapopulations, and even then by only 20%. The remaining four extant networks were expected to suffer extinction in 15-126 years. MVM was consistently estimated as approximately 5% of land area based on a sensitivity analysis of IFM parameters and was reduced only marginally (to approximately 4%) by modeling the potential impact of long-distance colonization over wider landscapes. The results suggest a widespread extinction debt among extant metapopulations of a declining species, necessitating conservation management or reserve designation even in apparent strongholds. For threatened species, metapopulation modeling is a potential means to identify landscapes near to extinction thresholds, to which conservation measures can be targeted for the best chance of success.     

Habitat-quality effects on metapopulation dynamics in greater white-toothed shrews, Crocidura russula

The effects of patch size and isolation on metapopulation dynamics have received wide empirical support and theoretical formalization. By contrast, the effects of patch quality seem largely underinvestigated, partly due to technical difficulties in properly assessing quality. Here we combine habitat-quality modeling with four years of demographic monitoring in a metapopulation of greater white-toothed shrews (Crocidura russula) to investigate the role of patch quality on metapopulation processes. Together, local patch quality and connectivity significantly enhanced local population sizes and occupancy rates (R2 = 14% and 19%, respectively). Accounting for the quality of patches connected to the focal one and acting as potential sources improved slightly the model explanatory power for local population sizes, pointing to significant source-sink dynamics. Local habitat quality, in interaction with connectivity, also increased colonization rate (R2 = 28%), suggesting the ability of immigrants to target high-quality patches. Overall, patterns were best explained when assuming a mean dispersal distance of 800 m, a realistic value for the species under study. Our results thus provide evidence that patch quality, in interaction with connectivity, may affect major demographic processes.