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.     

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.     

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.     

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.     

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.     

Mature and old-growth riparian forests: structure, dynamics, and effects on Adirondack stream habitats.

Riparian forests regulate linkages between terrestrial and aquatic ecosystems, yet relationships among riparian forest development, stand structure, and stream habitats are poorly understood in many temperate deciduous forest systems. Our research has (1) described structural attributes associated with old-growth riparian forests and (2) assessed linkages between these characteristics and in-stream habitat structure. The 19 study sites were located along predominantly first- and second-order streams in northern hardwood-conifer forests in the Adirondack Mountains of New York (U.S.A.). Sites were classified as mature forest (6 sites), mature with remnant old-growth trees (3 sites), and old-growth (10 sites). Forest-structure attributes were measured over stream channels and at varying distances from each bank. In-stream habitat features such as large woody debris (LWD), pools, and boulders were measured in each stream reach. Forest structure was examined in relation to stand age using multivariate techniques, ANOVA, and linear regression. We investigated linkages between forest structure and stream characteristics using similar methods, preceded by information-theoretic modeling (AIC). Old-growth riparian forest structure is more complex than that found in mature forests and exhibits significantly greater accumulations of aboveground tree biomass, both living and dead. In-stream LWD volumes were significantly (alpha = 0.05) greater at old-growth sites (200 m3/ha) compared to mature sites (34 m3/ha) and were strongly related to the basal area of adjacent forests. In-stream large-log densities correlated strongly with debris-dam densities. AIC models that included large-log density, debris-dam density, boulder density, and bankfull width had the most support for predicting pool density. There were higher proportions of LWD-formed pools relative to boulder-formed pools at old-growth sites as compared to mature sites. Old-growth riparian forests provide in-stream habitat features that have not been widely recognized in eastern North America, representing a potential benefit from late-successional riparian forest management and conservation. Riparian management practices (including buffer delineation and restorative silvicultural approaches) that emphasize development and maintenance of late-successional characteristics are recommended where the associated in-stream effects are desired.     

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.     

Assessing hypotheses about nesting site occupancy dynamics

Hypotheses about habitat selection developed in the evolutionary ecology framework assume that individuals, under some conditions, select breeding habitat based on expected fitness in different habitat. The relationship between habitat quality and fitness may be reflected by breeding success of individuals, which may in turn be used to assess habitat quality. Habitat quality may also be assessed via local density: if high-quality sites are preferentially used, high density may reflect high-quality habitat. Here we assessed whether site occupancy dynamics vary with site surrogates for habitat quality. We modeled nest site use probability in a seabird subcolony (the Black-legged Kittiwake, Rissa tridactyla) over a 20-year period. We estimated site persistence (an occupied site remains occupied from time t to t+1) and colonization through two subprocesses: first colonization (site creation at the timescale of the study) and recolonization (a site is colonized again after being deserted). Our model explicitly incorporated site-specific and neighboring breeding success and conspecific density in the neighborhood. Our results provided evidence that reproductively "successful" sites have a higher persistence probability than "unsuccessful" ones. Analyses of site fidelity in marked birds and of survival probability showed that high site persistence predominantly reflects site fidelity, not immediate colonization by new owners after emigration or death of previous owners. There is a negative quadratic relationship between local density and persistence probability. First colonization probability decreases with density, whereas recolonization probability is constant. This highlights the importance of distinguishing initial colonization and recolonization to understand site occupancy. All dynamics varied positively with neighboring breeding success. We found evidence of a positive interaction between site-specific and neighboring breeding success. We addressed local population dynamics using a site occupancy approach integrating hypotheses developed in behavioral ecology to account for individual decisions. This allows development of models of population and metapopulation dynamics that explicitly incorporate ecological and evolutionary processes.     

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.     

The importance of spatial autocorrelation,extent and resolution in predicting forest bird occurrence

Concerns about declines in forest biodiversity underscore the need for accurate estimates of the distribution and abundance of organisms at large scales and at resolutions that are fine enough to be appropriate for management. This paper addresses three major objectives: (i) to determine whether the resolution of typical air photo-derived forest inventory is sufficient for the accurate prediction of site occupancy by forest birds. We compared prediction success of habitat models using air photo variables to models with variables derived from finer resolution, ground-sampled vegetation plots. (ii) To test whether incorporating spatial autocorrelation into habitat models via autologistic regression increases prediction success. (iii) To determine whether landscape structure is an important factor in predicting bird distribution in forest-dominated landscapes. Models were tested locally (Greater Fundy Ecosystem [GFE]) using cross-validation, and regionally using an independent data set from an area located ca. 250 km to the northwest (Riley Brook [RB]). We found significant positive spatial autocorrelation in the residuals of at least one habitat model for 76% (16/21) of species examined. In these cases, the logistic regression assumption of spatially independent errors was violated. Logistic models that ignored spatial autocorrelation tended to overestimate habitat effects. Though overall prediction success was higher for autologistic models than logistic models in the GFE, the difference was only significantly improved for one species. Further, the inclusion of spatial covariates did little to improve model performance in the geographically discrete study area. For 62% (13/21) of species examined, landscape variables were significant predictors of forest bird occurrence even after statistically controlling for stand-level variability. However, broad spatial extents explained less variation than local factors. In the GFE, 76% (16/21) of air photo and 81% (17/21) of ground plot models were accurate enough to be of practical utility (AUC > 0.7). When applied to RB, both model types performed effectively for 55% (11/20) of the species examined. We did not detect an overall difference in prediction success between air photo and ground plot models in either study area. We conclude that air photo data are as effective as fine resolution vegetation data for predicting site occupancy for the majority of species in this study. These models will be of use to forest managers who are interested in mapping species distributions under various timber harvest scenarios, and to protected areas planners attempting to optimize reserve function.     

Local Population Dynamics in Metapopulation Models: Implications for Conservation

Abstract: Habitat fragmentation and the division of populations into spatially separated units have led to the increasing use of metapopulation models to characterize these populations. One prominent model that has served as a heuristic tool was introduced by Levins and is based on a collection of simplifying assumptions that exclude information on the dynamics and spatial distribution of local populations. Levins's and similar models predict the proportion of occupied habitat patches at equilibrium and the conditions needed to avoid total extinction. There are many obvious concerns about using such models, including how realistic alterations might change the predictions and whether occupancy has any relationship to population-level processes. Although many of the assumptions of these simple models are known to be unrealistic, we do not know how the assumptions affect model predictions. We simulated a metapopulation, and our results show that assumptions such as homogeneity of habitat patches, random migration among patches, equivalent extinction probabilities in all patches, and a large number of patches can lead to large overestimations of habitat occupancy. But when we explicitly modeled the underlying population dynamics within each patch, we found (1) that there was a strong correlation between proportion of occupied patches and total metapopulation size and (2) that the distribution of individuals among patches was relatively insensitive to model assumptions. Thus, our results show that although realistic modifications will change model predictions for occupancy, occupancy and population trends will be correlated. These correlations between occupancy and population size suggest that occupancy models may have some utility in conservation applications.     

Improving occupancy estimation when two types of observational error occur: non-detection and species misidentification

Efforts to draw inferences about species occurrence frequently account for false negatives, the common situation when individuals of a species are not detected even when a site is occupied. However, recent studies suggest the need to also deal with false positives, which occur when species are misidentified so that a species is recorded as detected when a site is unoccupied. Bias in estimators of occupancy, colonization, and extinction can be severe when false positives occur. Accordingly, we propose models that simultaneously account for both types of error. Our approach can be used to improve estimates of occupancy for study designs where a subset of detections is of a type or method for which false positives can be assumed to not occur. We illustrate properties of the estimators with simulations and data for three species of frogs. We show that models that account for possible misidentification have greater support (lower AIC for two species) and can yield substantially different occupancy estimates than those that do not. When the potential for misidentification exists, researchers should consider analytical techniques that can account for this source of error, such as those presented here.     

Community occupancy responses of small mammals to restoration treatments in ponderosa pine forests, northern Arizona, USA

In western North American conifer forests, wildfires are increasing in frequency and severity due to heavy fuel loads that have accumulated after a century of fire suppression. Forest restoration treatments (e.g., thinning and/or burning) are being designed and implemented at large spatial and temporal scales in an effort to reduce fire risk and restore forest structure and function. In ponderosa pine (Pinus ponderosa) forests, predominantly open forest structure and a frequent, low-severity fire regime constituted the evolutionary environment for wildlife that persisted for thousands of years. Small mammals are important in forest ecosystems as prey and in affecting primary production and decomposition. During 2006-2009, we trapped eight species of small mammals at 294 sites in northern Arizona and used occupancy modeling to determine community responses to thinning and habitat features. The most important covariates in predicting small mammal occupancy were understory vegetation cover, large snags, and treatment. Our analysis identified two generalist species found at relatively high occupancy rates across all sites, four open-forest species that responded positively to treatment, and two dense-forest species that responded negatively to treatment unless specific habitat features were retained. Our results indicate that all eight small mammal species can benefit from restoration treatments, particularly if aspects of their evolutionary environment (e.g., large trees, snags, woody debris) are restored. The occupancy modeling approach we used resulted in precise species-level estimates of occupancy in response to habitat attributes for a greater number of small mammal species than in other comparable studies. We recommend our approach for other studies faced with high variability and broad spatial and temporal scales in assessing impacts of treatments or habitat alteration on wildlife species. Moreover, since forest planning efforts are increasingly focusing on progressively larger treatment implementation, better and more efficiently obtained ecological information is needed to inform these efforts.     

Effects of landscape transformation on bird colonization and extinction patterns in a large‐scale,long‐term natural experiment

Conversion of agricultural land to forest plantations is a major driver of global change. Studies on the impact of forest plantations on biodiversity in plantations and in the surrounding native vegetation have been inconclusive. Consequently, it is not known how to best manage the extensive areas of the planet currently covered by plantations. We used a novel, long‐term (16 years) and large‐scale (30,000 ha) landscape transformation natural experiment (the Nanangroe experiment, Australia) to test the effects of land conversion on population dynamics of 64 bird species associated with woodland and forest. A unique aspect of our study is that we focused on the effects of plantations on birds in habitat patches within plantations. Our study design included 56 treatment sites (Eucalyptus patches where the surrounding matrix was converted from grazed land to pine plantations), 55 control sites (Eucalyptus patches surrounded by grazed land), and 20 matrix sites (sites within the pine plantations and grazed land). Bird populations were studied through point counts, and colonization and extinction patterns were inferred through multiple season occupancy models. Large‐scale pine plantation establishment affected the colonization or extinction patterns of 89% of studied species and thus led to a comprehensive turnover in bird communities inhabiting Eucalyptus patches embedded within the maturing plantations. Smaller bodied species appeared to respond positively to plantations (i.e., colonization increased and extirpation of these species decreased in patches surrounded by plantations) because they were able to use the newly created surrounding matrix. We found that the effects of forest plantations affected the majority of the bird community, and we believe these effects could lead to the artificial selection of one group of species at the expense of another.     

Conserving tigers in working landscapes

Tiger (Panthera tigris) conservation efforts in Asia are focused on protected areas embedded in human‐dominated landscapes. A system of protected areas is an effective conservation strategy for many endangered species if the network is large enough to support stable metapopulations. The long‐term conservation of tigers requires that the species be able to meet some of its life‐history needs beyond the boundaries of small protected areas and within the working landscape, including multiple‐use forests with logging and high human use. However, understanding of factors that promote or limit the occurrence of tigers in working landscapes is incomplete. We assessed the relative influence of protection status, prey occurrence, extent of grasslands, intensity of human use, and patch connectivity on tiger occurrence in the 5400 km² Central Terai Landscape of India, adjacent to Nepal. Two observer teams independently surveyed 1009 km of forest trails and water courses distributed across 60 166‐km² cells. In each cell, the teams recorded detection of tiger signs along evenly spaced trail segments. We used occupancy models that permitted multiscale analysis of spatially correlated data to estimate cell‐scale occupancy and segment‐scale habitat use by tigers as a function of management and environmental covariates. Prey availability and habitat quality, rather than protected‐area designation, influenced tiger occupancy. Tiger occupancy was low in some protected areas in India that were connected to extensive areas of tiger habitat in Nepal, which brings into question the efficacy of current protection and management strategies in both India and Nepal. At a finer spatial scale, tiger habitat use was high in trail segments associated with abundant prey and large grasslands, but it declined as human and livestock use increased. We speculate that riparian grasslands may provide tigers with critical refugia from human activity in the daytime and thereby promote tiger occurrence in some multiple‐use forests. Restrictions on human‐use in high‐quality tiger habitat in multiple‐use forests may complement existing protected areas and collectively promote the persistence of tiger populations in working landscapes.     

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.     

The Importance of Being Spatial (and Reserved): Assessing Northern Spotted Owl Habitat Relationships with Hierarchical Bayesian Models

Abstract:  Regional conservation planning increasingly draws on habitat suitability models to support decisions regarding land allocation and management. Nevertheless, statistical techniques commonly used for developing such models may give misleading results because they fail to account for 3 factors common in data sets of species distribution: spatial autocorrelation, the large number of sites where the species is absent (zero inflation), and uneven survey effort. We used spatial autoregressive models fit with Bayesian Markov Chain Monte Carlo techniques to assess the relationship between older coniferous forest and the abundance of Northern Spotted Owl nest and activity sites throughout the species' range. The spatial random-effect term incorporated in the autoregressive models successfully accounted for zero inflation and reduced the effect of survey bias on estimates of species–habitat associations. Our results support the hypothesis that the relationship between owl distribution and older forest varies with latitude. A quadratic relationship between owl abundance and older forest was evident in the southern portion of the range, and a pseudothreshold relationship was evident in the northern portion of the range. Our results suggest that proposed changes to the network of owl habitat reserves would reduce the proportion of the population protected by up to one-third, and that proposed guidelines for forest management within reserves underestimate the proportion of older forest associated with maximum owl abundance and inappropriately generalize threshold relationships among subregions. Bayesian spatial models can greatly enhance the utility of habitat analysis for conservation planning because they add the statistical flexibility necessary for analyzing regional survey data while retaining the interpretability of simpler models.     

A test of the substitution–habitat hypothesis in amphibians

Most examples that support the substitution‐habitat hypothesis (human‐made habitats act as substitutes of original habitat) deal with birds and mammals. We tested this hypothesis in 14 amphibians by using percentage occupancy as a proxy of habitat quality (i.e., higher occupancy percentages indicate higher quality). We classified water body types as original habitat (no or little human influence) depending on anatomical, behavioral, or physiological adaptations of each amphibian species. Ten species had relatively high probabilities (0.16–0.28) of occurrence in original habitat, moderate probability of occurrence in substitution habitats (0.11–0.14), and low probability of occurrence in refuge habitats (0.05–0.08). Thus, the substitution–habitat hypothesis only partially applies to amphibians because the low occupancy of refuges could be due to the negligible human persecution of this group (indicating good conservation status). However, low occupancy of refuges could also be due to low tolerance of refuge conditions, which could have led to selective extinction or colonization problems due to poor dispersal capabilities. That original habitats had the highest probabilities of occupancy suggests amphibians have a good conservation status in the region. They also appeared highly adaptable to anthropogenic substitution habitats.     

A robust-design formulation of the incidence function model of metapopulation dynamics applied to two species of rails

The incidence function model (IFM) uses area and connectivity to predict metapopulation dynamics. However, false absences and missing data can lead to underestimates of the number of sites contributing to connectivity, resulting in overestimates of dispersal ability and turnovers (extinctions plus colonizations). We extend estimation methods for the IFM by using a hierarchical Bayesian model to account both for false absences due to imperfect detection and for missing data due to sites not surveyed in some years. We compare parameter estimates, measures of metapopulation dynamics, and forecasts using stochastic patch occupancy models (SPOMs) among three IFM models: (1) a Bayesian formulation assuming no false absences and omitting site-year combinations with missing data; (2) a hierarchical Bayesian formulation assuming no false absences but incorporating missing data; and (3) a hierarchical Bayesian formulation allowing for imperfect detection and incorporating missing data. We fit the models to multiyear data sets of occupancy for two bird species that differ in body size and presumed dispersal ability but inhabit the same network of sites: the small Black Rail (Laterallus jamaicensis) and the medium-sized Virginia Rail (Rallus limicola). Incorporating missing data affected colonization parameters and led to lower estimates of dispersal ability for the Black Rail. Detection rates were high for the Black Rail in most years but moderate for the Virginia Rail. Incorporating imperfect detection resulted in higher occupancy and lower turnover rates for both species, with largest effects for the Virginia Rail. Forecasts using SPOMs were sensitive to both missing data and false absences; persistence in models assuming no false absences was more optimistic than from robust models. Our results suggest that incorporating false absences and missing data into the IFM can improve (1) estimates of dispersal ability and the effect of connectivity on colonization, (2) the scaling of extinction risk with patch area, and (3) forecasts of occupancy and turnover rates.