Rapid Increases and Time‐Lagged Declines in Amphibian Occupancy after Wildfire

Climate change is expected to increase the frequency and severity of drought and wildfire. Aquatic and moisture‐sensitive species, such as amphibians, may be particularly vulnerable to these modified disturbance regimes because large wildfires often occur during extended droughts and thus may compound environmental threats. However, understanding of the effects of wildfires on amphibians in forests with long fire‐return intervals is limited. Numerous stand‐replacing wildfires have occurred since 1988 in Glacier National Park (Montana, U.S.A.), where we have conducted long‐term monitoring of amphibians. We measured responses of 3 amphibian species to fires of different sizes, severity, and age in a small geographic area with uniform management. We used data from wetlands associated with 6 wildfires that burned between 1988 and 2003 to evaluate whether burn extent and severity and interactions between wildfire and wetland isolation affected the distribution of breeding populations. We measured responses with models that accounted for imperfect detection to estimate occupancy during prefire (0–4 years) and different postfire recovery periods. For the long‐toed salamander (Ambystoma macrodactylum) and Columbia spotted frog (Rana luteiventris), occupancy was not affected for 6 years after wildfire. But 7–21 years after wildfire, occupancy for both species decreased ≥25% in areas where >50% of the forest within 500 m of wetlands burned. In contrast, occupancy of the boreal toad (Anaxyrus boreas) tripled in the 3 years after low‐elevation forests burned. This increase in occupancy was followed by a gradual decline. Our results show that accounting for magnitude of change and time lags is critical to understanding population dynamics of amphibians after large disturbances. Our results also inform understanding of the potential threat of increases in wildfire frequency or severity to amphibians in the region. Incrementos Rápidos y Declinaciones Desfasadas en la Ocupación de Anfibios Después de un Incendio     

Using multilevel spatial models to understand salamander site occupancy patterns after wildfire

Studies of the distribution of elusive forest wildlife have suffered from the confounding of true presence with the uncertainty of detection. Occupancy modeling, which incorporates probabilities of species detection conditional on presence, is an emerging approach for reducing observation bias. However, the current likelihood modeling framework is restrictive for handling unexplained sources of variation in the response that may occur when there are dependence structures such as smaller sampling units that are nested within larger sampling units. We used multilevel Bayesian occupancy modeling to handle dependence structures and to partition sources of variation in occupancy of sites by terrestrial salamanders (family Plethodontidae) within and surrounding an earlier wildfire in western Oregon, USA. Comparison of model fit favored a spatial N-mixture model that accounted for variation in salamander abundance over models that were based on binary detection/non-detection data. Though catch per unit effort was higher in burned areas than unburned, there was strong support that this pattern was due to a higher probability of capture for individuals in burned plots. Within the burn, the odds of capturing an individual given it was present were 2.06 times the odds outside the burn, reflecting reduced complexity of ground cover in the burn. Ther was weak support that true occupancy was lower within the burned area. While the odds of occupancy in the burn were 0.49 times the odds outside the burn among the five species, the magnitude of variation attributed to the burn was small in comparison to variation attributed to other landscape variables and to unexplained, spatially autocorrelated random variation. While ordinary occupancy models may separate the biological pattern of interest from variation in detection probability when all sources of variation are known, the addition of random effects structures for unexplained sources of variation in occupancy and detection probability may often more appropriately represent levels of uncertainty.     

Effects of Urbanization on Occupancy of Stream Salamanders

Abstract: Urban development is the most common form of land conversion in the United States. Using a before–after control‐impact study design, we investigated the effects of urbanization on larval and adult stages of southern two‐lined salamanders (Eurycea cirrigera) and northern dusky salamanders (Desmognathus fuscus). Over 5 years, we estimated changes in occupancy and probabilities of colonization and survival in 13 stream catchments after urbanization and in 17 catchments that were not urbanized. We also examined effects of proportion of urbanized area in a catchment and distance of the salamander population to the nearest stream on probabilities of colonization and survival. Before urbanization, adult and larval stages of the two salamander species occupied nearly all surveyed streams, with occupancy estimates ranging from 1.0 to 0.78. Four years after urbanization mean occupancy of larval and adult two‐lined salamanders had decreased from 0.87 and 0.78 to 0.57 and 0.39, respectively. Estimates of mean occupancy of larval northern dusky salamanders decreased from 1.0 to 0.57 in urban streams 4 years after urbanization; however, adult northern dusky salamander occupancy remained close to 1.0 in urban streams over 5 years. Occupancy estimates in control streams were similar for each species and stage over 5 years. Urbanization was associated with decreases in survival probabilities of adult and larval two‐lined salamanders and decreases in colonization probabilities of larval dusky salamanders. Nevertheless, proportion of impervious surface and distance to nearest stream had little effect on probabilities of survival and colonization. Our results imply that in the evaluation of the effects of urbanization on species, such as amphibians, with complex life cycles, consideration of the effects of urbanization on both adult and larval stages is required.     

Spatial Patterns of Breeding Success of Grizzly Bears Derived from Hierarchical Multistate Models

Conservation programs often manage populations indirectly through the landscapes in which they live. Empirically, linking reproductive success with landscape structure and anthropogenic change is a first step in understanding and managing the spatial mechanisms that affect reproduction, but this link is not sufficiently informed by data. Hierarchical multistate occupancy models can forge these links by estimating spatial patterns of reproductive success across landscapes. To illustrate, we surveyed the occurrence of grizzly bears (Ursus arctos) in the Canadian Rocky Mountains Alberta, Canada. We deployed camera traps for 6 weeks at 54 surveys sites in different types of land cover. We used hierarchical multistate occupancy models to estimate probability of detection, grizzly bear occupancy, and probability of reproductive success at each site. Grizzly bear occupancy varied among cover types and was greater in herbaceous alpine ecotones than in low‐elevation wetlands or mid‐elevation conifer forests. The conditional probability of reproductive success given grizzly bear occupancy was 30% (SE = 0.14). Grizzly bears with cubs had a higher probability of detection than grizzly bears without cubs, but sites were correctly classified as being occupied by breeding females 49% of the time based on raw data and thus would have been underestimated by half. Repeated surveys and multistate modeling reduced the probability of misclassifying sites occupied by breeders as unoccupied to <2%. The probability of breeding grizzly bear occupancy varied across the landscape. Those patches with highest probabilities of breeding occupancy—herbaceous alpine ecotones—were small and highly dispersed and are projected to shrink as treelines advance due to climate warming. Understanding spatial correlates in breeding distribution is a key requirement for species conservation in the face of climate change and can help identify priorities for landscape management and protection. Patrones Espaciales del Éxito Reproductivo de Osos Pardos, Derivados de Modelos Jerárquicos Multi‐Estado     

Human influence on California fire regimes.

Periodic wildfire maintains the integrity and species composition of many ecosystems, including the mediterranean-climate shrublands of California. However, human activities alter natural fire regimes, which can lead to cascading ecological effects. Increased human ignitions at the wildland-urban interface (WUI) have recently gained attention, but fire activity and risk are typically estimated using only biophysical variables. Our goal was to determine how humans influence fire in California and to examine whether this influence was linear, by relating contemporary (2000) and historic (1960-2000) fire data to both human and biophysical variables. Data for the human variables included fine-resolution maps of the WUI produced using housing density and land cover data. Interface WUI, where development abuts wildland vegetation, was differentiated from intermix WUI, where development intermingles with wildland vegetation. Additional explanatory variables included distance to WUI, population density, road density, vegetation type, and ecoregion. All data were summarized at the county level and analyzed using bivariate and multiple regression methods. We found highly significant relationships between humans and fire on the contemporary landscape, and our models explained fire frequency (R2 = 0.72) better than area burned (R2 = 0.50). Population density, intermix WUI, and distance to WUI explained the most variability in fire frequency, suggesting that the spatial pattern of development may be an important variable to consider when estimating fire risk. We found nonlinear effects such that fire frequency and area burned were highest at intermediate levels of human activity, but declined beyond certain thresholds. Human activities also explained change in fire frequency and area burned (1960-2000), but our models had greater explanatory power during the years 1960-1980, when there was more dramatic change in fire frequency. Understanding wildfire as a function of the spatial arrangement of ignitions and fuels on the landscape, in addition to nonlinear relationships, will be important to fire managers and conservation planners because fire risk may be related to specific levels of housing density that can be accounted for in land use planning. With more fires occurring in close proximity to human infrastructure, there may also be devastating ecological impacts if development continues to grow farther into wildland vegetation.     

A Bayesian state-space formulation of dynamic occupancy models

Species occurrence and its dynamic components, extinction and colonization probabilities, are focal quantities in biogeography and metapopulation biology, and for species conservation assessments. It has been increasingly appreciated that these parameters must be estimated separately from detection probability to avoid the biases induced by non-detection error. Hence, there is now considerable theoretical and practical interest in dynamic occupancy models that contain explicit representations of metapopulation dynamics such as extinction, colonization, and turnover as well as growth rates. We describe a hierarchical parameterization of these models that is analogous to the state-space formulation of models in time series, where the model is represented by two components, one for the partially observable occupancy process and another for the observations conditional on that process. This parameterization naturally allows estimation of all parameters of the conventional approach to occupancy models, but in addition, yields great flexibility and extensibility, e.g., to modeling heterogeneity or latent structure in model parameters. We also highlight the important distinction between population and finite sample inference; the latter yields much more precise estimates for the particular sample at hand. Finite sample estimates can easily be obtained using the state-space representation of the model but are difficult to obtain under the conventional approach of likelihood-based estimation. We use R and WinBUGS to apply the model to two examples. In a standard analysis for the European Crossbill in a large Swiss monitoring program, we fit a model with year-specific parameters. Estimates of the dynamic parameters varied greatly among years, highlighting the irruptive population dynamics of that species. In the second example, we analyze route occupancy of Cerulean Warblers in the North American Breeding Bird Survey (BBS) using a model allowing for site-specific heterogeneity in model parameters. The results indicate relatively low turnover and a stable distribution of Cerulean Warblers which is in contrast to analyses of counts of individuals from the same survey that indicate important declines. This discrepancy illustrates the inertia in occupancy relative to actual abundance. Furthermore, the model reveals a declining patch survival probability, and increasing turnover, toward the edge of the range of the species, which is consistent with metapopulation perspectives on the genesis of range edges. Given detection/non-detection data, dynamic occupancy models as described here have considerable potential for the study of distributions and range dynamics.     

Wildfire smoke and health impacts: A closer look at fire attributes and their marginal effects

Existing studies on the economic impact of wildfire smoke have focused on single fire events or entire seasons without considering the marginal effect of daily fire progression on downwind communities. Neither approach allows for an examination of the impact of even the most basic fire attributes, such as distance and fuel type, on air quality and health outcomes. Improved knowledge of these effects can provide important guidance for efficient wildfire management strategies. This study aims to bridge this gap using detailed information on 24 large-scale wildfires that sent smoke plumes to the Reno/Sparks area of Northern Nevada over a 4-year period. We relate the daily acreage burned by these fires to daily data on air pollutants and local hospital admissions. Using information on medical expenses, we compute the per-acre health cost of wildfires of different attributes. We find that patient counts can be causally linked to fires as far as 200–300 miles from the impact area. As expected, the marginal impact per acre burned generally diminishes with distance and for fires with lighter fuel loads. Our results also highlight the importance of allowing for temporal lags between fire occurrence and pollutant levels.     

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.     

Use of Multispecies Occupancy Models to Evaluate the Response of Bird Communities to Forest Degradation Associated with Logging

Forest degradation is arguably the greatest threat to biodiversity, ecosystem services, and rural livelihoods. Therefore, increasing understanding of how organisms respond to degradation is essential for management and conservation planning. We were motivated by the need for rapid and practical analytical tools to assess the influence of management and degradation on biodiversity and system state in areas subject to rapid environmental change. We compared bird community composition and size in managed (ejido, i.e., communally owned lands) and unmanaged (national park) forests in the Sierra Tarahumara region, Mexico, using multispecies occupancy models and data from a 2‐year breeding bird survey. Unmanaged sites had on average higher species occupancy and richness than managed sites. Most species were present in low numbers as indicated by lower values of detection and occupancy associated with logging‐induced degradation. Less than 10% of species had occupancy probabilities >0.5, and degradation had no positive effects on occupancy. The estimated metacommunity size of 125 exceeded previous estimates for the region, and sites with mature trees and uneven‐aged forest stand characteristics contained the highest species richness. Higher estimation uncertainty and decreases in richness and occupancy for all species, including habitat generalists, were associated with degraded young, even‐aged stands. Our findings show that multispecies occupancy methods provide tractable measures of biodiversity and system state and valuable decision support for landholders and managers. These techniques can be used to rapidly address gaps in biodiversity information, threats to biodiversity, and vulnerabilities of species of interest on a landscape level, even in degraded or fast‐changing environments. Moreover, such tools may be particularly relevant in the assessment of species richness and distribution in a wide array of habitats. Uso de Modelos de Ocupación para Múltiples Especies para Evaluar la Respuesta de las Comunidades de Aves a la Degradación de Bosques Asociada con la Tala     

Postfire Logging in Riparian Areas

Abstract:  We reviewed the behavior of wildfire in riparian zones, primarily in the western United States, and the potential ecological consequences of postfire logging. Fire behavior in riparian zones is complex, but many aquatic and riparian organisms exhibit a suite of adaptations that allow relatively rapid recovery after fire. Unless constrained by other factors, fish tend to rebound relatively quickly, usually within a decade after a wildfire. Additionally, fire and subsequent erosion events contribute wood and coarse sediment that can create and maintain productive aquatic habitats over time. The potential effects of postfire logging in riparian areas depend on the landscape context and disturbance history of a site; however, available evidence suggests two key management implications: (1) fire in riparian areas creates conditions that may not require intervention to sustain the long-term productivity of the aquatic network and (2) protection of burned riparian areas gives priority to what is left rather than what is removed. Research is needed to determine how postfire logging in riparian areas has affected the spread of invasive species and the vulnerability of upland forests to insect and disease outbreaks and how postfire logging will affect the frequency and behavior of future fires. The effectiveness of using postfire logging to restore desired riparian structure and function is therefore unproven, but such projects are gaining interest with the departure of forest conditions from those that existed prior to timber harvest, fire suppression, and climate change. In the absence of reliable information about the potential consequence of postfire timber harvest, we conclude that providing postfire riparian zones with the same environmental protections they received before they burned is justified ecologically. Without a commitment to monitor management experiments, the effects of postfire riparian logging will remain unknown and highly contentious.     

Effects of river-flow regulation on anuran occupancy and abundance in riparian zones

The natural flow regimes of rivers worldwide have been heavily altered through anthropogenic activities, and dams in particular have a pervasive effect on riverine ecosystems. Flow-regulation effects of dams negatively affect species diversity and abundance of a variety of aquatic animals, including invertebrates and fishes. However, the effects on semiaquatic animals are relatively unknown. We conducted anuran calling surveys at 42 study locations along the Broad and Pacolet Rivers in South Carolina to address the potential effects of flow regulation by damming on anuran occupancy and abundance. We estimated occupancy and abundance with Program PRESENCE. Models incorporated distance upstream and downstream from the nearest dam as covariates and urbanization pressure as an alternative stressor. Distance from dam was associated with occupancy of 2 of the 9 anuran species in our analyses and with abundance of 6 species. In all cases, distance downstream from nearest dam was a better predictor of occupancy and abundance than distance upstream from nearest dam. For all but one species, distance downstream from nearest dam was positively correlated with both occupancy and abundance. Reduced occupancy and abundance of anurans likely resulted from downstream alterations in flow regime associated with damming, which can lead to reduced area of riparian wetlands that serve as anuran breeding habitat. Our results showed that damming has a strong negative effect on multiple anuran species across large spatial extents and suggest that flow regulation can affect semiaquatic animals occupying riparian zones.     

Effects of Traffic Noise on Occupancy Patterns of Forest Birds

Abstract: Noise may drive changes in the composition and abundance of animals that communicate vocally. Traffic produces low‐frequency noise (<3 kHz) that can mask acoustic signals broadcast within the same frequency range. We evaluated whether birds that sing within the frequency range of traffic noise are affected by acoustic masking (i.e., increased background noise levels at the same frequency of vocalizations reduce detection of vocalization) and are less abundant in areas where traffic noise is loud (44–57 dB). We estimated occupancy, the expected probability that a given site is occupied by a species, and detection probabilities of eight forest‐breeding birds in areas with and without traffic noise as a function of noise and three measures of habitat quality: percent forest cover, distance from plot center to the edge of forest, and the number of standing dead trees or snags. For the two species that vocalize at the lowest peak frequency (the frequency with the most energy) and the lowest overall frequency (Yellow‐billed Cuckoo [Coccyzus americanus] and White‐breasted Nuthatch [Sitta carolinensis]), the presence of traffic noise explained the greatest proportion of variance in occupancy, and these species were 10 times less likely to be found in noisy than in quiet plots. For species that had only portions of their vocalizations overlapped by traffic noise, either forest cover or distance to forest edge explained more variation in occupancy than noise or no single variable explained occupancy. Our results suggest that the effects of traffic noise may be especially pronounced for species that vocalize at low frequencies.     

Modeling mammal response to fire based on species’ traits

Fire has shaped ecological communities worldwide for millennia, but impacts of fire on individual species are often poorly understood. We performed a meta-analysis to predict which traits, habitat, or study variables and fire characteristics affect how mammal species respond to fire. We modeled effect sizes of measures of population abundance or occupancy as a function of various combinations of these traits and variables with phylogenetic least squares regression. Nine of 115 modeled species (7.83%) returned statistically significant effect sizes, suggesting most mammals are resilient to fire. The top-ranked model predicted a negative impact of fire on species with lower reproductive rates, regardless of fire type (estimate = –0.68), a positive impact of burrowing in prescribed fires (estimate = 1.46) but not wildfires, and a positive impact of average fire return interval for wildfires (estimate = 0.93) but not prescribed fires. If a species’ International Union for Conservation of Nature Red List assessment includes fire as a known or possible threat, the species was predicted to respond negatively to wildfire relative to prescribed fire (estimate = –2.84). These findings provide evidence of experts’ abilities to predict whether fire is a threat to a mammal species and the ability of managers to meet the needs of fire-threatened species through prescribed fire. Where empirical data are lacking, our methods provide a basis for predicting mammal responses to fire and thus can guide conservation actions or interventions in species or communities.     

Cicada emergence in Southwestern riparian forest: influences of wildfire and vegetation composition.

Annually emerging cicadas are a numerically and ecologically dominant species in Southwestern riparian forests. Humans have altered disturbance regimes that structure these forests such that floods are less common and wildfires occur more frequently than was historically the case. Impacts of these changes on primary consumers such as riparian cicadas are unknown. Because cicadas are consumed by a variety of animal species, disturbances that alter timing of their emergence or abundance could have consequences for species at higher trophic levels. We trapped emerging cicadas (Tibicen dealbatus) in burned and unburned riparian forest plots along the Middle Rio Grande in central New Mexico (USA) to determine effects of wildfire and vegetation structure on their density and phenology. We measured vegetation variables and soil temperature at cicada traps and related these variables to variation in emergence density and phenology. We also experimentally heated soil under emergence traps to examine the relationship between soil temperature and emergence phenology. Emergence density was similar in wildfire and unburned plots, though emergence date averaged earlier in wildfire plots and experimentally heated traps. We identified models containing cottonwood proximity (distance from the nearest cottonwood tree) and cottonwood canopy coverage as the most parsimonious explanations of emergence density at each trap. Model selection results were consistent with the literature and field observations that showed that cottonwood trees are an essential resource for T. dealbatus. Cottonwood canopy was also correlated with low soil temperatures, which are associated with later emergence dates. Failure of cottonwoods to reestablish following wildfire could result in cicadas emerging at lower densities and at earlier dates. For cicadas to emerge at densities and times that provide the greatest benefits to birds and other riparian-obligate secondary consumers, riparian forests should be protected from fire, and native vegetation in wildfire sites should be restored.     

Effects of Postfire Salvage Logging on Deadwood‐Associated Beetles

Abstract: In Canada and the United States pressure to recoup financial costs of wildfire by harvesting burned timber is increasing, despite insufficient understanding of the ecological consequences of postfire salvage logging. We compared the species richness and composition of deadwood‐associated beetle assemblages among undisturbed, recently burned, logged, and salvage‐logged, boreal, mixed‐wood stands. Species richness was lowest in salvage‐logged stands, largely due to a negative effect of harvesting on the occurrence of wood‐ and bark‐boring species. In comparison with undisturbed stands, the combination of wildfire and logging in salvage‐logged stands had a greater effect on species composition than either disturbance alone. Strong differences in species composition among stand treatments were linked to differences in quantity and quality (e.g., decay stage) of coarse woody debris. We found that the effects of wildfire and logging on deadwood‐associated beetles were synergistic, such that the effects of postfire salvage logging could not be predicted reliably on the basis of data on either disturbance alone. Thus, increases in salvage logging of burned forests may have serious negative consequences for deadwood‐associated beetles and their ecological functions in early postfire successional forests.     

Metapopulation dynamics in the butterfly Hipparchia semele changed decades before occupancy declined in The Netherlands

The survival of many species in human-dominated, fragmented landscapes depends on metapopulation dynamics, i.e., on a dynamic equilibrium of extinctions and colonizations in patches of suitable habitat. To understand and predict distributional changes, knowledge of these dynamics can be essential, and for this, metapopulation studies are preferably based on long-time-series data from many sites. Alas, such data are very scarce. An alternative is to use opportunistic data (i.e., collected without applying standardized field methods), but these data suffer from large variations in field methods and search intensity between sites and years. Dynamic site-occupancy models offer a general approach to adjust for variable survey effort. These models extend classical metapopulation models to account for imperfect detection of species and yield estimates of the probabilities of occupancy, colonization, and survival of species at sites. By accounting for detection, they fully correct for among-year variability in search effort. As an illustration, we fitted a dynamic site-occupancy model to 60 years of presence-absence data (more precisely, detection-nondetection) of the heathland butterfly Hipparchia semele in The Netherlands. Detection records were obtained from a database containing volunteer-based data from 1950-2009, and nondetection records were deduced from database records of other butterfly species. Our model revealed that metapopulation dynamics of Hipparchia had changed decades before the species' distribution began to contract. Colonization probability had already started to decline from 1950 onward, but this was counterbalanced by an increase in the survival of existing populations, the result of which was a stable distribution. Only from 1990 onward was survival not sufficient to compensate for the further decrease of colonization, and occupancy started to decline. Hence, it appears that factors acting many decades ago triggered a change in the metapopulation dynamics of this species, which ultimately led to a severe decline in occupancy that only became apparent much later. Our study emphasizes the importance of knowledge of changes in survival and colonization of species in modern landscapes over a very long time scale. It also demonstrates the power of site-occupancy modeling to obtain important population dynamics information from databases containing opportunistic sighting records.     

Patch-Occupancy Modeling as a Method for Monitoring Changes in Forest Floristics: a Case Study in Southeastern Australia

Abstract:  The ability to monitor changes in biodiversity is fundamental to demonstrating sustainable management practices of natural resources. Disturbance studies generally focus on responses at the plot scale, whereas landscape-scale responses are directly relevant to the development of sustainable forest management. Modeling changes in occupancy is one way to monitor landscape-scale responses. We used understory vegetation data collected over 16 years from a long-term study site in southeastern Australia. The site was subject to timber harvesting and frequent prescribed burning. We used occupancy models to examine the impacts of these disturbances on the distribution of 50 species of plants during the study. Timber harvesting influenced the distribution of 9 species, but these effects of harvesting were generally lost within 14 years. Repeated prescribed fire affected 22 species, but the heterogeneity of the burns reduced the predicted negative effects. Twenty-two species decreased over time independent of treatment, and only 5 species increased over time. These changes probably represent a natural response to a wildfire that occurred in 1973, 13 years before the study began. Occupancy modeling is a useful and flexible technique for analyzing monitoring data and it may also be suitable for inclusion within an adaptive-management framework for forest management.     

Rapid assessment of postfire plant invasions in coniferous forests of the western United States.

Fire is a natural part of most forest ecosystems in the western United States, but its effects on nonnative plant invasion have only recently been studied. Also, forest managers are engaging in fuel reduction projects to lessen fire severity, often without considering potential negative ecological consequences such as nonnative plant species introductions. Increased availability of light, nutrients, and bare ground have all been associated with high-severity fires and fuel treatments and are known to aid in the establishment of nonnative plant species. We use vegetation and environmental data collected after wildfires at seven sites in coniferous forests in the western United States to study responses of nonnative plants to wildfire. We compared burned vs. unburned plots and plots treated with mechanical thinning and/or prescribed burning vs. untreated plots for nonnative plant species richness and cover and used correlation analyses to infer the effect of abiotic site conditions on invasibility. Wildfire was responsible for significant increases in nonnative species richness and cover, and a significant decrease in native cover. Mechanical thinning and prescribed fire fuel treatments were associated with significant changes in plant species composition at some sites. Treatment effects across sites were minimal and inconclusive due to significant site and site x treatment interaction effects caused by variation between sites including differences in treatment and fire severities and initial conditions (e.g., nonnative species sources). We used canonical correspondence analysis (CCA) to determine what combinations of environmental variables best explained patterns of nonnative plant species richness and cover. Variables related to fire severity, soil nutrients, and elevation explained most of the variation in species composition. Nonnative species were generally associated with sites with higher fire severity, elevation, percentage of bare ground, and lower soil nutrient levels and lower canopy cover. Early assessments of postfire stand conditions can guide rapid responses to nonnative plant invasions.     

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.     

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.