Search

Context Recent increases in ecological disturbances driven by climate change and our expanding human footprint make it challenging for natural resource managers to keep apprised of current conditions and adjust management plans accordingly. To effectively conserve species in highly dynamic landscapes requires more timely habitat monitoring and a more responsive adaptive management cycle. Objectives We introduce a framework to automatically monitor and assess species habitats over a range of spatial and temporal scales. We then apply this framework by developing an automated habitat monitoring system for the Mexican spotted owl (MSO) in Arizona and New Mexico, USA, that will be linked to federal agency adaptive management plans. Methods We automated the process of monitoring and assessing trends in MSO habitat on an annual schedule using the Google Earth Engine cloud-based spatial analysis platform and dynamic data repository. We ran this system retrospectively on historical data to monitor MSO habitat from 1986 to 2020. Results The automated habitat monitoring system provided a 35-year MSO habitat time series with high accuracy. Widespread habitat gains and losses occurred every year, underscoring the need for continuous monitoring and the benefits of an automated workflow. Conclusions Automated habitat monitoring linked to adaptive management holds great promise in helping managers track the impacts of recent disturbances and adjust plans to meet goals even in increasingly dynamic landscapes. In a companion paper, Jones et al. (2023) demonstrate the utility of this approach by analyzing our MSO habitat time series to assess trends, drivers of change, and management implications.

Context Understanding habitat dynamics is essential for effective conservation as landscapes rapidly change. In a companion paper in this issue, Shirk et al. (2023) introduced an automated habitat monitoring system using Google Earth Engine and applied this framework to develop a dynamic model of Mexican spotted owl (Strix occidentalis lucida) habitat across the southwestern US from 1986 to 2020. Objectives We explored the application of this dynamic model of Mexican spotted owl habitat in the context of the species’ ecology. Methods We evaluated environmental correlates of Mexican spotted owl habitat, assessed potential spatial non-stationarity in habitat selection, estimated long-term trends in habitat by quantifying changes in habitat amount and quality between 1986 and 2020, and evaluated the extent to which habitat changes over the past 35 years have been driven by wildfire. Results Topography and climate appeared to outweigh reflectance-based (vegetation) metrics in describing Mexican spotted owl habitat and habitat selection was non-stationary across modeling sub-regions. Total habitat area for Mexican spotted owls declined by ~ 21% since 1986 (0.6% annually), but trends varied spatially and some even reversed over the past decade. Wildfire was responsible for between 8 and 35% of total habitat loss, depending on the sub-region considered. Conclusions The automated habitat monitoring system allowed trend estimation and accurate assessment of current habitat status for Mexican spotted owls; maps were accurate, spatially detailed, and current. The ability to continually produce accurate maps for large land areas for threatened species such as the Mexican spotted owl facilitates science-based land management on public lands in the southwestern US.

We conducted a range-wide investigation of the dynamics of site-level reproductive rate of northern spotted owls using survey data from 11 study areas across the subspecies geographic range collected during 1993-2018. Our analytical approach accounted for imperfect detection of owl pairs and misclassification of successful reproduction (i.e., at least one young fledged) and contributed further insights into northern spotted owl population ecology and dynamics. Both nondetection and state misclassification were important, especially because factors affecting these sources of error also affected focal ecological parameters. Annual probabilities of site occupancy were greatest at sites with successful reproduction in the previous year and lowest for sites not occupied by a pair in the previous year. Site-specific occupancy transition probabilities declined over time and were negatively affected by barred owl presence. Overall, the site-specific probability of successful reproduction showed substantial year-to-year fluctuations and was similar for occupied sites that did or did not experience successful reproduction the previous year. Site-specific probabilities for successful reproduction were very small for sites that were unoccupied the previous year. Barred owl presence negatively affected the probability of successful reproduction by northern spotted owls in Washington and California, as predicted, but the effect in Oregon was mixed. The proportions of sites occupied by northern spotted owl pairs showed steep, near-monotonic declines over the study period, with all study areas showing the lowest observed levels of occupancy to date. If trends continue it is likely that northern spotted owls will become extirpated throughout large portions of their range in the coming decades.

Background The northern spotted owl (Strix occidentalis caurina) is an Endangered Species Act-listed subspecies that requires coniferous forests with structurally complex and closed-canopy old-growth characteristics for nesting. With climate change, large wildfires are expected to become more common within the subspecies’ range and an increasing threat to these types of forests. Understanding fire severity patterns related to suitable nesting forest will be important to inform forest management that affects conservation and recovery. We examined the relationship between fire severity and suitable nesting forest in 472 large wildfires (> 200 ha) that occurred in the northern spotted owl range during 1987–2017. We mapped fire severities (unburned-low, moderate, high) within each fire using relative differenced normalized burn ratios and quantified differences in severity between pre-fire suitable nesting forest (edge and interior) and non-nesting forest. We also quantified these relationships within areas of three fire regimes (low severity, very frequent; mixed severity, frequent; high severity, infrequent). Results Averaged over all fires, the interior nesting forest burned at lower severity than edge or non-nesting forest. These relationships were consistent within the low severity, very frequent, and mixed severity, frequent fire regime areas. All forest types burned at similar severity within the high severity, infrequent fire regime. During two of the most active wildfire years that also had the largest wildfires occurring in rare and extreme weather conditions, we found a bimodal distribution of fire severity in all forest types. In those years, a higher amount—and proportion—of all forest types burned at high severity. Over the 30-year study, we found a strong positive trend in the proportion of wildfires that burned at high severity in the non-nesting forests, but not in the suitable nesting forest types. Conclusions Under most wildfire conditions, the microclimate of interior patches of suitable nesting forests likely mitigated fire severity and thus functioned as fire refugia (i.e., burning at lower severity than the surrounding landscape). With changing climate, the future of interior forest as fire refugia is unknown, but trends suggest older forests can dampen the effect of increased wildfire activity and be an important component of landscapes with fire resiliency.

Owls are important avian predators in forested systems, but little is known about landscape use by most forest‐adapted owl species in environments impacted by mixed‐severity wildfire. To better understand species‐specific patterns of post‐wildfire landscape use within an owl guild, we used passive acoustic monitoring using autonomous recording units. The technology is effective for multi‐species surveys, especially if some species are rare, nocturnal, or difficult to detect by traditional means. In 2017, we surveyed the interior and adjacent unburned areas of a 10,700‐ha mixed‐severity wildfire that burned in 2015 in southwest Oregon. We used occupancy modeling to identify patterns of landscape use by five species of forest owls: barred owls (Strix varia), great horned owls (Bubo virginianus), western screech‐owls (Megascops kennicottii), northern pygmy‐owls (Glaucidium gnoma), and northern saw‐whet owls (Aegolius acadicus). Our results showed a positive relationship between increasing fire severity and probability of use by western screech‐owls and a similar but somewhat weaker relationship for northern pygmy‐owls. Barred owls were rarely detected in severely burned areas and their use decreased with increased fire severity. We observed generally low landscape use for great horned owls, which decreased with increased fire severity and at higher elevations. Thus, four out of the five species appeared to use recently burned forests at different levels, with only northern saw‐whet owls showing near‐complete avoidance of the burned area. These findings increase our understanding of the basic ecology of each species and highlight the varied use of burned areas within this community. These previously undocumented patterns of landscape use in burned landscapes should provide insights to managers and policymakers in the Pacific Northwest as climate shifts, and fires may increase in size, frequency, and severity.

Significance Invasive species can cause extinctions of native species and widespread biodiversity loss. Invader removal is a common management response, but the use of long-term field experiments to characterize effectiveness of removals in benefitting impacted native species is rare. We used a large-scale removal experiment to investigate the demographic response of a threatened native species, the northern spotted owl, to removal of an invasive competitor species, the barred owl. Removal of barred owls had a strong, positive effect on survival of spotted owls, which arrested long-term population declines of spotted owls. The results demonstrate that the long-term persistence of spotted owls will depend heavily on reducing the negative impacts of barred owls while simultaneously addressing other threats, such as habitat loss., Changes in the distribution and abundance of invasive species can have far-reaching ecological consequences. Programs to control invaders are common but gauging the effectiveness of such programs using carefully controlled, large-scale field experiments is rare, especially at higher trophic levels. Experimental manipulations coupled with long-term demographic monitoring can reveal the mechanistic underpinnings of interspecific competition among apex predators and suggest mitigation options for invasive species. We used a large-scale before–after control–impact removal experiment to investigate the effects of an invasive competitor, the barred owl (Strix varia), on the population dynamics of an iconic old-forest native species, the northern spotted owl (Strix occidentalis caurina). Removal of barred owls had a strong, positive effect on survival of sympatric spotted owls and a weaker but positive effect on spotted owl dispersal and recruitment. After removals, the estimated mean annual rate of population change for spotted owls stabilized in areas with removals (0.2% decline per year), but continued to decline sharply in areas without removals (12.1% decline per year). The results demonstrated that the most substantial changes in population dynamics of northern spotted owls over the past two decades were associated with the invasion, population expansion, and subsequent removal of barred owls. Our study provides experimental evidence of the demographic consequences of competitive release, where a threatened avian predator was freed from restrictions imposed on its population dynamics with the removal of a competitively dominant invasive species.

Background Many mammalian species have experienced range contractions. Following a reduction in distribution that has resulted in apparently small and disjunct populations, the Humboldt marten (Martes caurina humboldtensis) was recently designated as federally Threatened and state Endangered. This subspecies of Pacific marten occurring in coastal Oregon and northern California, also known as coastal martens, appear unlike martens that occur in snow-associated regions in that vegetation associations appear to differ widely between Humboldt marten populations. We expected current distributions represent realized niches, but estimating factors associated with long-term occurrence was challenging for this rare and little-known species. Here, we assessed the predicted contemporary distribution of Humboldt martens and interpret our findings as hypotheses correlated with the subspecies’ niche to inform strategic conservation actions. Methods We modeled Humboldt marten distribution using a maximum entropy (Maxent) approach. We spatially-thinned 10,229 marten locations collected from 1996–2020 by applying a minimum distance of 500-m between locations, resulting in 384 locations used to assess correlations of marten occurrence with biotic and abiotic variables. We independently optimized the spatial scale of each variable and focused development of model variables on biotic associations (e.g., hypothesized relationships with forest conditions), given that abiotic factors such as precipitation are largely static and not alterable within a management context. Results Humboldt marten locations were positively associated with increased shrub cover (salal (Gautheria shallon)), mast producing trees (e.g., tanoak, Notholithocarpus densiflorus), increased pine (Pinus sp.) proportion of total basal area, annual precipitation at home-range spatial scales, low and high amounts of canopy cover and slope, and cooler August temperatures. Unlike other recent literature, we found little evidence that Humboldt martens were associated with old-growth structural indices. This case study provides an example of how limited information on rare or lesser-known species can lead to differing interpretations, emphasizing the need for study-level replication in ecology. Humboldt marten conservation would benefit from continued survey effort to clarify range extent, population sizes, and fine-scale habitat use.

The US Endangered Species Act has enabled species conservation but has differentially impacted fire management and rare bird conservation in the southern and western US. In the South, prescribed fire and restoration-based forest thinning are commonly used to conserve the endangered red-cockaded woodpecker (Picoides borealis; RCW), whereas in the West, land managers continue to suppress fire across the diverse habitats of the northern, Californian, and Mexican spotted owls (Strix occidentalis subspecies; SO). Although the habitat needs of the RCW and SO are not identical, substantial portions of both species’ ranges have historically been exposed to relatively frequent, low-to moderate-intensity fires. Active management with fire and thinning has benefited the RCW but proves challenging in the western US. We suggest the western US could benefit from the adoption of a similar innovative approach through policy, public–private partnerships, and complementarity of endangered species management with multiple objectives. These changes would likely balance long-term goals of SO conservation and enhance forest resilience.

Background The northern spotted owl (Strix occidentalis caurina) is an Endangered Species Act-listed subspecies that requires forests with old-growth characteristics for nesting. With climate change, large, severe wildfires are expected to be more common and an increasing threat to spotted owl persistence. Understanding fire severity patterns related to nesting forest can be valuable for forest management that supports conservation and recovery, especially if nesting forest functions as fire refugia (i.e., lower fire severity than surrounding landscape). We examined the relationship between fire severity and nesting forests in 472 large wildfires (> 200 ha) that occurred rangewide during 1987–2017. We mapped fire severities (unburned-low, moderate, high) within each fire using relative difference normalized burn ratios and quantified differences in severity between pre-fire nesting forest (edge and interior) and non-nesting forest. We also quantified these relationships within areas of three fire regimes (low severity, very frequent; mixed severity, frequent; high severity, infrequent). Results Averaged over all fires, the interior nesting forest burned at lower severity than edge or non-nesting forest. These relationships were consistent within the low severity, very frequent and mixed severity, frequent fire regime areas. All forest types burned at similar severity within the high severity, infrequent fire regime. During two of the most active wildfire years that also had the largest wildfires occurring in rare and extreme weather conditions, we found a bimodal distribution of fire severity in all forest types. In those years, a higher amount—and proportion—of all forest types burned at high severity. Over the duration of the study, we found a strong positive trend in the proportion of wildfires that burned at high severity in the non-nesting forests, but not in the two nesting forest types. Conclusions Under most wildfire conditions, the microclimate of interior patches of nesting forests likely mitigated fire severity and thus functioned as fire refugia. With changing climates, the future of interior forest as fire refugia is unknown, but trends suggest these older forests can dampen the effect of increased wildfire activity and thus an important component of landscape plans focused on fire resiliency.

BackgroundA suite of mammalian species have experienced range contractions following European settlement and post-settlement development of the North American continent. For example, while North American martens (American marten, Martes americana; Pacific marten, M. caurina) generally have a broad range across northern latitudes, local populations have experienced substantial reductions in distribution and some extant populations are small and geographically isolated. The Humboldt marten (M. c. humboldtensis), a subspecies of Pacific marten that occurs in coastal Oregon and northern California, was recently designated as federally threatened in part due to its reduced distribution. To inform strategic conservation actions, we assessed Humboldt marten occurrence by compiling all known records from their range.MethodsWe compiled Humboldt marten locations since their rediscover to present (1,692 marten locations, 1996-2020). We spatially-thinned locations to 500-m to assess correlations with variables across contemporary Humboldt marten distribution (n=384). Using maximum entropy modeling (Maxent), we created distribution models with variables optimized for spatial scale; pre-selected scales were associated with marten ecology (50 to 1170 m radius). Marten locations were most correlated with abiotic factors (e.g., precipitation), which are unalterable and therefore uninformative within the context of restoration or management actions. Thus, we created variables to focus on hypothesized marten habitat relationships, including understory conditions such as predicted suitability of shrub species.ResultsHumboldt marten locations were positively associated with increased shrub cover (salal (Gautheria shallon), mast producing trees), increased pine (Pinus sp) overstory cover and precipitation at home-range spatial scales, areas with low and high amounts of canopy cover and slope, and cooler August temperatures. Unlike other recent literature on the species, we found little evidence that Humboldt marten locations were associated with old growth structural indices, perhaps because of a potential mismatch in the association between this index and shrub cover. As with any species distribution model, there were gaps in predicted distribution where Humboldt martens have been located during more recent surveys, for instance the southeastern portion of Oregon’s coast range. Conservation efforts and our assessment of potential risks to Humboldt marten populations would benefit from additional information on range extent, population sizes, and fine-scale habitat use. Like many rare and lesser-known species, this case study provides an example of how limited information can provide differing interpretations, emphasizing the need for study-level replication in ecology.

Frogs dependent on lotic environments are sensitive to disturbances that alter the hydrology (e.g., water impoundments), substrate (e.g., debris torrents), and riparian vegetation (e.g., wildfires) of river ecosystems. Although rivers are often very dynamic, disturbances can push environmental baselines outside of narrowly defined ecological tolerances under which a species evolved. Short-lived lotic-dependent organisms, restricted to movements within the water or the riparian corridor, are at risk of local extirpations owing to such disturbances if they fragment and isolate affected populations from recolonizing source populations. In Oregon, USA, the foothill yellow-legged frog (Rana boylii) is at its northernmost range margin and has experienced an approximately 41% range contraction compared to their historical distribution. To inform conservation and management, we used species distribution models to identify environmentally suitable watersheds based on intrinsic baseline environmental variables, and then examined potential effects of human-caused alterations to rivers, including splash dams used to ferry timber downstream prior to 1957, large water impoundments, and adjacency to agricultural croplands. We used machine-learning in program Maxent and three different river layers that varied in extent and location of mapped rivers but contained distinct information to produce species distribution models which we then combined into a single ensemble model. Stream order, annual precipitation, and precipitation frequency were the highest ranked baseline environmental variables in most models. Watersheds with highly suitable baseline conditions in our ensemble model were negatively correlated with anthropogenic disturbances to rivers. Foothill yellow-legged frogs appeared to be sensitive to human-caused disturbances to rivers, perhaps indicative of their narrow ecological tolerance to in-river conditions. We do not anticipate variables in our model to change much through time. Rather, for conservation we identified potential legacy (spash dams) and ongoing human-caused disturbances that are more likely to change conditions for the species in the short- and long-term.

Western North American forest ecosystems are experiencing rapid changes in disturbance regimes because of climate change and land use legacies (Littell et al. 2018). In many of these forests, the accumulation of surface and ladder fuels from a century of fire suppression, coupled with a warming and drying climate, has led to increases in the number of large fires (Westerling 2016) and the proportion of areas burning at higher severity (Safford and Stevens 2017, Singleton et al. 2018). While the annual area burned by fire is still below historical levels (Taylor et al. 2016), some forest types in the west are burning at higher severities when compared to pre-European settlement periods (Mallek et al. 2013, Safford and Stevens 2017). As such, they face an increased risk of conversion to non-forest ecosystems (e.g., shrublands, non-native grasslands) following large, severe fires because of compromised seed sources, post-fire soil erosion and loss, high-severity re-burn, and climatic thresholds (Coppoletta et al. 2016, Stevens et al. 2017, Rissman et al. 2018, Shive et al. 2018, Wood and Jones 2019). Restoration methods such as mechanical thinning and prescribed and managed wildland fire that reduce accumulated surface and ladder fuels (e.g., removal of smalland medium-sized trees, especially non-fire adapted species) may reduce the spatial extent of severe fires and increase forest resilience to fire in a changing climate (Agee and Skinner 2005, Stephens et al. 2013, Hessburg et al. 2016, Tubbesing et al. 2019) and, in doing so, promote key ecosystem services (Hurteau et al. 2014, Kelsey et al. 2017, Wood and Jones 2019).

Author(s): Krawchuk, Meg A; Meigs, Garrett W; Cartwright, Jennifer M; Coop, Jonathan D; Davis, Raymond; Holz, Andres; Kolden, Crystal; Meddens, Arjan JH

Passive acoustic monitoring using autonomous recording units (ARUs) is a fast-growing area of wildlife research especially for rare, cryptic species that vocalize. Northern Spotted Owl (Strix occidentalis caurina) populations have been monitored since the mid-1980s using mark–recapture methods. To evaluate an alternative survey method, we used ARUs to detect calls of Northern Spotted Owls and Barred Owls (S. varia), a congener that has expanded its range into the Pacific Northwest and threatens Northern Spotted Owl persistence. We set ARUs at 30 500-ha hexagons (150 ARU stations) with recent Northern Spotted Owl activity and high Barred Owl density within Northern Spotted Owl demographic study areas in Oregon and Washington, and set ARUs to record continuously each night from March to July, 2017. We reviewed spectrograms (visual representations of sound) and tagged target vocalizations to extract calls from ~160,000 hr of recordings. Even in a study area with low occupancy rates on historical territories (Washington’s Olympic Peninsula), the probability of detecting a Northern Spotted Owl when it was present in a hexagon exceeded 0.95 after 3 weeks of recording. Environmental noise, mainly from rain, wind, and streams, decreased detection probabilities for both species over all study areas. Using demographic information about known Northern Spotted Owls, we found that weekly detection probabilities of Northern Spotted Owls were higher when ARUs were closer to known nests and activity centers and when owls were paired, suggesting passive acoustic data alone could help locate Northern Spotted Owl pairs on the landscape. These results demonstrate that ARUs can effectively detect Northern Spotted Owls when they are present, even in a landscape with high Barred Owl density, thereby facilitating the use of passive, occupancy-based study designs to monitor Northern Spotted Owl populations.

The northern spotted owl (Strix occidentalis caurina) inhabits older coniferous forests in the Pacific Northwest and has been at the center of forest management issues in this region. The immediate threats to this federally listed species include habitat loss and competition with barred owls (Strix varia), which invaded from eastern North America. We conducted a prospective meta-analysis to assess population trends and factors affecting those trends in northern spotted owls using 26 years of survey and capture-recapture data from 11 study areas across the owls' geographic range to analyze demographic traits, rates of population change, and occupancy parameters for spotted owl territories. We found that northern spotted owl populations experienced significant declines of 6–9% annually on 6 study areas and 2–5% annually on 5 other study areas. Annual declines translated to ≤35% of the populations remaining on 7 study areas since 1995. Barred owl presence on spotted owl territories was the primary factor negatively affecting apparent survival, recruitment, and ultimately, rates of population change. Analysis of spotted and barred owl detections in an occupancy framework corroborated the capture-recapture analyses with barred owl presence increasing territorial extinction and decreasing territorial colonization of spotted owls. While landscape habitat components reduced the effect of barred owls on these rates of decline, they did not reverse the negative trend. Our analyses indicated that northern spotted owl populations potentially face extirpation if the negative effects of barred owls are not ameliorated while maintaining northern spotted owl habitat across their range.

Contraction of native old forest can limit occurrence of old forest associated species, especially species with limited vagility. Patterns of size and distribution of remaining patches of old forest along with forest disturbance and what replaces old forest can influence whether species adapt or perish after forest loss. The arboreal red tree vole (Arborimus longicaudus) is a small arvicoline rodent that is associated with old coniferous forest and typically emigrates short distances. Since 1911, old forest (≥80 years old) in the northern half of the Oregon Coast Range has been reduced by >80%, primarily due to large stand replacing wildfires, timber harvest, and subsequent conversion to young forest (

We modeled the normal fire environment for occurrence of large forest wildfires (>40 ha) for the Pacific Northwest Region of the United States. Large forest wildfire occurrence data from the recent climate normal period (1971–2000) was used as the response variable and fire season precipitation, maximum temperature, slope, and elevation were used as predictor variables. A projection of our model onto the 2001–2030 climate normal period showed strong agreement between model predictions and the area of forest burned by large wildfires from 2001 to 2015 (independent fire data). We then used downscaled climate projections for two greenhouse gas concentration scenarios and over 30 climate models to project changes in environmental suitability for large forest fires over the 21st century. Results indicated an increasing proportion of forested area with fire environments more suitable for the occurrence of large wildfires over the next century for all ecoregions but less pronounced for the Coast Range and Puget Lowlands. The largest increases occurred on federal lands, while private and state lands showed less. We calculated fire rotation periods for the recent historical and current climate and examined the relative differences between them and our modeled large wildfire suitability classes. By the end of the century, the models predicted shorter fire rotation periods, with cooler/moister forests experiencing larger magnitudes of change than warmer/drier forests. Modeling products, including a set of time series maps, can provide forest resource managers, fire protection agencies, and policy-makers empirical estimates of how much and where climate change might affect the geographic distribution of large wildfires and effect fire rotations.

The northern spotted owl (Strix occidentalis caurina) was listed as threatened under the U.S. Endangered Species Act (ESA) in 1990. We applied modern spatial conservation theory and models to evaluate several candidate critical habitat networks, and sought an efficient conservation solution that encompassed the highest value lands for spotted owl recovery rather than maximizing the total area of potential critical habitat. We created a map of relative habitat suitability, which served as input to the spatial conservation prioritization program Zonation. We used the spatially-explicit individual-based population model HexSim to estimate and compare simulated spotted owl population outcomes among a suite of candidate critical habitat networks that varied in size and spatial arrangement under alternative scenarios of future habitat suitability and barred owl (S. varia) effects. We evaluated simulated spotted owl population outcomes, including total population size, and extinction and quasi-extinction likelihoods for 108 combinations of candidate critical habitat networks by habitat change by barred owl scenarios, both range-wide and within 11 distinct portions of the owl’s range. Barred owl encounter rates and the amount and suitability of habitat had substantial effects on simulated spotted owl populations. When barred owl encounter rates were high, changes in the amount and suitability of habitat had minimal impacts on population performance. Under lowered barred owl encounter rates, candidate critical habitat networks that included most existing high suitability habitat supported a high likelihood of long-term population persistence. Barred owls are currently the primary driving force behind poor population performance of NSOs; however, our models demonstrated that a sufficient area of high suitability habitat remains essential for recovery when effects of barred owls can be reduced. The modeling approach we employed is sufficiently flexible to incorporate new information about spotted owls as it becomes available and could likely be applied to conservation planning for other species.

In the Pacific Northwest of the United States, 20th century timber harvesting resulted in major declines in area of forests supporting large live and dead trees (i.e., snags), that are not only key habitat elements for many wildlife species but also critical components of ecosystem function. Regional forest management guidance, such as the Northwest Forest Plan (1994) and Eastside Screens (1995), may aim to conserve and foster the development of late-successional old-growth forests, characterized by large live and dead trees. Satellite remote sensing supports regional monitoring efforts of these habitat characteristics, but managers may require additional guidance in order to leverage these data for large landscape or regional assessments. In this study, our objectives were to assess long-term (historical vs. contemporary) and short-term (1993–2017) changes in lands supporting large live trees and snags across 10 wildlife habitat types (WHTs) – disparate vegetation conditions that support significantly different wildlife communities – in Oregon and Washington, USA. We generated 30-m, annual maps of large live trees (>50 cm, >75 cm, and >100 cm diameter) and snags (>25 cm and > 50 cm diameter) based on the gradient nearest neighbor (GNN) imputation method. GNN integrates Landsat satellite imagery, geospatial climatic and topographic data, and USDA Forest Service Forest Inventory and Analysis data to predict forest attributes for all forested lands in the study area. GNN classification accuracy was poor to good for large live trees (Cohen’s kappa = 0.2 – 0.6) and poor to fair for snags (Cohen’s kappa = 0.1 – 0.3) in most WHTs, though performance was substantially lower in drier WHTs where large live trees and snags were rare. Our results highlighted long-term reductions in forest supporting large live trees and snags from historical to contemporary times, especially in wetter, more productive WHTs. In contrast, we observed short-term (1993–2017) increases in areas supporting large live trees and snags. Federal forests were both more similar to reference conditions and exhibited greater recent increases in areas supporting large live trees compared to nonfederal lands. Thus, Oregon and Washington have lost a substantial proportion of forests containing large live trees and snags and recent recruitment of these trees at regional scales is a slow process primarily occurring on federal lands. However, detecting such changes through current Landsat satellite mapping technologies remains challenging, highlighting the need for new mapping methods to aid in future management.

Occupancy modeling is a widely used analytical technique for assessing species distributions and range dynamics. However, occupancy analyses ignore variation in abundances of occupied sites, even though site abundances affect many of the parameters being estimated (e.g., extinction, colonization, detection probability). We introduce a new model (“dynamic N-occupancy”) capable of providing accurate estimates of local abundance, population gains (reproduction/immigration), and apparent survival probabilities while accounting for imperfect detection using only detection/nondetection data. Our model utilizes heterogeneity in detection based on variations in site abundances to estimate latent demographic rates via a dynamic N-mixture modeling framework. We validate our model using simulations across a wide range of values and examine the data requirements, including the number of years and survey sites needed, for unbiased and precise estimation of parameters. We apply our model to estimate spatio-temporal heterogeneity in abundances of barred owls (Strix varia) within a recently invaded region in Oregon (USA). Estimates of apparent survival and population gains are consistent with those from a nearby radio-tracking study and elucidate how barred owl abundances have increased dramatically over time. The dynamic N-occupancy model greatly improves inferences on individual-level population processes from occupancy data by explicitly modeling the latent population structure. This article is protected by copyright. All rights reserved.

Conservation planning for at-risk species requires understanding of where species are likely to occur, how many individuals are likely to be supported on a given landscape, and the ability to monitor those changes through time. We developed a distribution model for northern spotted owls that incorporates both habitat suitability and probability of territory occupancy while accounting for interspecies competition. We developed range-wide habitat suitability maps for two time periods (1993 and 2012) for northern spotted owls that accounted for regional differences in habitat use and home range size. We used these maps for a long-term demographic monitoring study area to assess habitat change and estimate the number of potential territories based on available habitat for both time periods. We adjusted the number of potential territories using known occupancy rates to estimate owl densities for both time periods. We evaluated our range-wide habitat suitability model using independent survey data. Our range-wide habitat maps predicted areas suitable for territorial spotted owl presence well. On the demographic study area, the amount of habitat declined 19.7% between 1993 and 2012, while our estimate of the habitat-based carrying capacity declined from 150 to 146 territories. Estimated number of occupied territories declined from 94 to 57. Conservation and recovery of at-risk species depends on understanding how habitat changes over time in response to factors such as wildfire, climate change, biological invasions, and interspecies competition, and how these changes influence species distribution. We demonstrate a model-based approach that provides an effective planning tool.

Slow ecological processes challenge conservation. Short-term variability can obscure the importance of slower processes that may ultimately determine the state of a system. Furthermore, management actions with slow responses can be hard to justify. One response to slow processes is to explicitly concentrate analysis on state dynamics. Here, we focus on identifying drivers of Northern Spotted Owl (Strix occidentalis caurina) territorial occupancy dynamics across 11 study areas spanning their geographic range and forecasting response to potential management actions. Competition with Barred Owls (Strix varia) has increased Spotted Owl territory extinction probabilities across all study areas and driven recent declines in Spotted Owl populations. Without management intervention, the Northern Spotted Owl subspecies will be extirpated from parts of its current range within decades. In the short term, Barred Owl removal can be effective. Over longer time spans, however, maintaining or improving habitat conditions can help promote the persistence of northern spotted owl populations. In most study areas, habitat effects on expected Northern Spotted Owl territorial occupancy are actually greater than the effects of competition from Barred Owls. This study suggests how intensive management actions (removal of a competitor) with rapid results can complement a slower management action (i.e., promoting forest succession).

Landscape genetics investigations examine how the availability and configuration of habitat influence genetic structure of plants and animals. We used landscape genetics to evaluate the role that forest connectivity plays in determining genetic structure of the federally-threatened Northern Spotted Owl (Strix occidentalis caurina) using genotypes of 339 Northern Spotted Owls obtained for 10 microsatellite loci. Spatial clustering analyses identified a distinct genetic cluster at the southern extent of the region examined. This cluster could not be linked to landscape connectivity patterns and suggested that post-Pleistocene processes were involved with its development rather than contemporary landscape configuration. We also compared matrices of pairwise inter-individual genetic distances with resistance distances derived from a circuit-theory based framework. Resistance distances were obtained for an idealized raster map that reflected continuous unimpeded dispersal habitat across the landscape along with five empirically-derived raster maps reflecting the 1870's, 1940's, 1986, 1994, and 2012. Resistance distances from the idealized map served as surrogates for linear geographic distances. Relative to idealized conditions, resistance distances were ~250% higher in the 1940's and ~200% higher from 1986 onward. Resistance distances from the 1870's were ~40% higher than idealized conditions. Inter-individual genetic distances were most highly correlated with resistance distances from the idealized map rather than any of the empirical maps. Two hypotheses explain our results. First, our results may reflect temporal lags between the onset of large-scale habitat alterations and their novel effects on genetic structure in long-lived species such as Northern Spotted Owls. Second, because Northern Spotted Owls disperse over long distances, our results may indicate that forest habitat has never been sufficiently fragmented to the point where connectivity was disrupted. The second hypothesis could indicate that forest management practices mandated by the Northwest Forest Plan succeeded with one of its primary goals. However, our results do not represent a complete portrayal of the status of Northern Spotted Owls given detection of significant population declines and bottlenecks in other studies. Future investigations based on computer simulations may help distinguish between hypotheses.

Estimates of species' vital rates and an understanding of the factors affecting those parameters over time and space can provide crucial information for management and conservation. We used mark-recapture, reproductive output, and territory occupancy data collected during 1985-2013 to evaluate population processes of Northern Spotted Owls (Strix occidentalis caurina) in 11 study areas in Washington, Oregon, and northern California, USA. We estimated apparent survival, fecundity, recruitment, rate of population change, and local extinction and colonization rates, and investigated relationships between these parameters and the amount of suitable habitat, local and regional variation in meteorological conditions, and competition with Barred Owls (Strix varia). Data were analyzed for each area separately and in a meta-analysis of all areas combined, following a strict protocol for data collection, preparation, and analysis. We used mixed effects linear models for analyses of fecundity, Cormack-Jolly-Seber open population models for analyses of apparent annual survival (phi), and a reparameterization of the Jolly-Seber capture-recapture model (i.e. reverse Jolly-Seber; RJS) to estimate annual rates of population change (lambda(RJS)) and recruitment. We also modeled territory occupancy dynamics of Northern Spotted Owls and Barred Owls in each study area using 2-species occupancy models. Estimated mean annual rates of population change (lambda) suggested that Spotted Owl populations declined from 1.2% to 8.4% per year depending on the study area. The weighted mean estimate of lambda for all study areas was 0.962 (+/- 0.019 SE; 95% CI: 0.925-0.999), indicating an estimated range-wide decline of 3.8% per year from 1985 to 2013. Variation in recruitment rates across the range of the Spotted Owl was best explained by an interaction between total winter precipitation and mean minimum winter temperature. Thus, recruitment rates were highest when both total precipitation (29 cm) and minimum winter temperature (-9.5 degrees C) were lowest. Barred Owl presence was associated with increased local extinction rates of Spotted Owl pairs for all 11 study areas. Habitat covariates were related to extinction rates for Spotted Owl pairs in 8 of 11 study areas, and a greater amount of suitable owl habitat was generally associated with decreased extinction rates. We observed negative effects of Barred Owl presence on colonization rates of Spotted Owl pairs in 5 of 11 study areas. The total amount of suitable Spotted Owl habitat was positively associated with colonization rates in 5 areas, and more habitat disturbance was associated with lower colonization rates in 2 areas. We observed strong declines in derived estimates of occupancy in all study areas. Mean fecundity of females was highest for adults (0.309 +/- 0.027 SE), intermediate for 2-yr-olds (0.179 +/- 0.040 SE), and lowest for 1-yr-olds (0.065 +/- 0.022 SE). The presence of Barred Owls and habitat covariates explained little of the temporal variation in fecundity in most study areas. Climate covariates occurred in competitive fecundity models in 8 of 11 study areas, but support for these relationships was generally weak. The fecundity meta-analysis resulted in 6 competitive models, all of which included the additive effects of geographic region and annual time variation. The 2 top-ranked models also weakly supported the additive negative effects of the amount of suitable core area habitat, Barred Owl presence, and the amount of edge habitat on fecundity. We found strong support for a negative effect of Barred Owl presence on apparent survival of Spotted Owls in 10 of 11 study areas, but found few strong effects of habitat on survival at the study area scale. Climate covariates occurred in top or competitive survival models for 10 of 11 study areas, and in most cases the relationships were as predicted; however, there was little consistency among areas regarding the relative importance of specific climate covariates. In contrast, meta-analysis results suggested that Spotted Owl survival was higher across all study areas when the Pacific Decadal Oscillation (PDO) was in a warming phase and the Southern Oscillation Index (SOI) was negative, with a strongly negative SOI indicative of El Nino events. The best model that included the Barred Owl covariate (BO) was ranked 4th and also included the PDO covariate, but the BO effect was strongly negative. Our results indicated that Northern Spotted Owl populations were declining throughout the range of the subspecies and that annual rates of decline were accelerating in many areas. We observed strong evidence that Barred Owls negatively affected Spotted Owl populations, primarily by decreasing apparent survival and increasing local territory extinction rates. However, the amount of suitable owl habitat, local weather, and regional climatic patterns also were related to survival, occupancy (via colonization rate), recruitment, and, to a lesser extent, fecundity, although there was inconsistency in regard to which covariates were important for particular demographic parameters or across study areas. In the study areas where habitat was an important source of variation for Spotted Owl demographics, vital rates were generally positively associated with a greater amount of suitable owl habitat. However, Barred Owl densities may now be high enough across the range of the Northern Spotted Owl that, despite the continued management and conservation of suitable owl habitat on federal lands, the long-term prognosis for the persistence of Northern Spotted Owls may be in question without additional management intervention. Based on our study, the removal of Barred Owls from the Green Diamond Resources (GDR) study area had rapid, positive effects on Northern Spotted Owl survival and the rate of population change, supporting the hypothesis that, along with habitat conservation and management, Barred Owl removal may be able to slow or reverse Northern Spotted Owl population declines on at least a localized scale. USDA Forest ServiceUnited States Department of Agriculture (USDA)United States Forest Service; USDI Bureau of Land Management; USDI National Park Service; Green Diamond Resource Company; Plum Creek Timber Company; Louisiana Pacific Lumber Company; Hancock Forest Management; Hoopa Tribe; USDI Bureau of Land Management via a Cost Reimbursable Research Agreement with Oregon State University Funding for demographic studies of Northern Spotted Owls on federal lands was provided primarily by the USDA Forest Service, USDI Bureau of Land Management, and USDI National Park Service. Funding for studies on nonfederal lands came from the Green Diamond Resource Company, Plum Creek Timber Company, Louisiana Pacific Lumber Company, Hancock Forest Management, and the Hoopa Tribe. Funding for the workshop was provided by the USDA Forest Service and USDI Bureau of Land Management via a Cost Reimbursable Research Agreement with Oregon State University. The U.S. Geological Survey and USDA Forest Service required approval of the final manuscript before publication, but none of the funding agencies associated with this meta-analysis had any influence on the content of the submitted or published manuscript. Any use of trade, firm, or product names is for descriptive purposes only and does not imply endorsement by the U.S. Government. Ethics statement: All data from study areas in which K.M.D., E.D.F., and A.B.F. were principal investigators (CLE, OLY, COA, HJA, TYE, CAS, and NWC) were collected following protocols approved under animal care and use permits overseen by Oregon State University's and Colorado State University's Institutional Animal Care and Use Committees. These protocols were followed and all relevant state and federal permits were acquired and maintained for all study areas included in this meta-analysis. Public domain – authored by a U.S. government employee

Accuracy assessments of remote sensing products are necessary for identifying map strengths and weaknesses in scientific and management applications. However, not all accuracy assessments are created equal. Motivated by a recent study published in Forest Ecology and Management (Volume 342, pages 8–20), we explored the potential limitations of accuracy assessments related to characteristics of the field data: sampling bias and spatial resolution. The authors of the previous paper used data from variable radius plots near northern spotted owl nest sites to assess the predictive accuracy of gradient nearest neighbor (GNN) maps in portions of Oregon and Washington, USA. The field plots used for accuracy assessment (1) potentially biased the accuracy assessment toward older forests and (2) examined accuracy at finer scales than the imputation map predictions under consideration. To examine both the impacts of bias and scale in accuracy assessment, we assessed the predictive accuracy of GNN maps in western and southern Oregon. We found correlation coefficients between predicted (900 m2) and observed forest attributes for small plots (506 m2) were consistently lower than accuracy assessments using larger plots (4048 m2). Similarly, correlation coefficients based only on field plots near nest sites were lower than correlations based on all field plots. These results imply that sampling bias and small plot areas result in accuracy assessments that underestimate map predictive performance. In particular, assessing accuracy at spatial scales below the resolution of the map products are overly pessimistic (i.e., low correlation coefficients). While accuracy assessment is important, care needs to be taken to ensure that the sampling design for field data does not limit inference on map accuracy.

article i nfo Wildlifehabitatmapping has evolved at a rapidpace over thelastfew decades. Beginningwith simple, oftensub- jective, hand-drawn maps, habitat mapping now involves complex species distribution models (SDMs) using mapped predictor variables derived from remotely sensed data. For species that inhabit large geographic areas, remote sensing technology is often essential for producing range wide maps. Habitat monitoring for northern spotted owls (Strix occidentalis caurina), whose geographic covers about 23 million ha, is based on SDMs that use Landsat Thematic Mapper imagery to create forest vegetation data layers using gradient nearest neighbor (GNN) methods. Vegetation data layers derived from GNN are modeled relationships between forest inventory plot data, climate and topographic data, and the spectral signatures acquired by the satellite. When used as pre- dictor variables for SDMs, there is some transference of the GNN modeling error to the final habitat map. Recent increasesinthe use of light detection and ranging (lidar) data, coupled with the need to produce spatially accurate and detailed forest vegetation maps have spurred interest in its use for SDMs and habitat mapping. In- stead of modeling predictor variables from remotely sensed spectral data, lidar provides direct measurements of vegetationheightforuseinSDMs.WeexpectaSDMhabitatmapproducedfromdirectlymeasuredpredictorvar- iables to be more accurate than one produced from modeled predictors. We used maximum entropy (Maxent) SDM modeling software to compare predictive performance and esti- mates of habitat area between Landsat-based and lidar-based northern spotted owl SDMs and habitat maps. We explored the differences and similarities between these maps, and to a pre-existing aerial photo- interpreted habitat map produced by local wildlife biologists. The lidar-based map had the highest predictive performance based on 10 bootstrapped replicate models (AUC = 0.809 ± 0.011), but the performance of the Landsat-based map was within acceptable limits (AUC = 0.717 ± 0.021). As is common with photo- interpreted maps, there was no accuracy assessment available for comparison. The photo-interpreted map pro- duced the highest and lowest estimates of habitat area, depending on which habitat classes were included (nesting, roosting, and foraging habitat = 9962 ha, nesting habitat only = 6036 ha). The Landsat-based map produced an estimate of habitat area that was within this range (95% CI: 6679-9592 ha), while the lidar-based map produced an area estimate similar to what was interpreted by local wildlife biologists as nesting (i.e., high quality) habitat using aerial imagery (95% CI: 5453-7216). Confidence intervals of habitat area estimates from the SDMs based on Landsat and lidar overlapped. Weconcludedthatboth Landsat- and lidar-based SDMs produced reasonablemaps and areaestimatesfor north- ern spotted owl habitat within the study area. The lidar-based map was more precise and spatially similar to whatlocal wildlife biologistsconsideredspottedowl nesting habitat.The Landsat-basedmap provideda lesspre- cisespatial representation of habitatwithin therelatively small geographic confines ofthe study area,but habitat area estimates were similar to both the photo-interpreted and lidar-based maps.

Pacific Northwest moist coniferous forests provide a wide array of globally important goods and services, including water, carbon sequestration, wood products, fish and wildlife habitat, cultural values, and world-class recreation. These forests are owned and managed by a mix of public, private, and tribal landowners (plates 6, 7), however, who often have different forest-management objectives. Overall, this diverse landownership provides a highly variable forest landscape with forest-management objectives ranging from intensive management on industrial forestlands, to longer rotations on state and tribal lands, to an emphasis on preservation and restoration of late-successional forests to support endangered species and water quality on federal lands. In this chapter, we synthesize some of the objectives of different landowners in the region and the potential opportunities and challenges of integrating goods and services (ecosystem services) into forest management. We show how broad assessment of ecosystem services can be used to plan management activities and to evaluate trade-offs of managing public and private lands to provide a suite of goods and services.

The scope and extent of past natural disturbances and human-derived changes to the forest landscape often provide the historical context for management but are often insufficiently accounted for in forest planning. In particular, static components of many management plans are not easily adapted to unforeseen system dynamics. For example, when the Northwest Forest Plan was designed in 1993, the inherently dynamic nature of the forest ecosystem and landscape was acknowledged, but there was a general lack of scientific information about the ecological processes that would shape forests of the future. The expectation was that both management and natural disturbances would influence change in the forested landscape, but how management would then adapt to these altered conditions was not clear. At the time, climate change was not well understood and was just beginning to be discussed in relation to forests.

The role of competition in structuring biotic communities at fine spatial scales is well known from detailed process-based studies. Our understanding of competition's importance at broader scales is less resolved and mainly based on static species distribution maps. Here, we bridge this gap by examining the joint occupancy dynamics of an invading species (Barred Owl, Strix varia) and a resident species (Northern Spotted Owl, Strix occidentalis caurina) in a 1000-km study area over a 22-year period. Past studies of these competitors have focused on the dynamics of one species at a time, hindering efforts to parse out the roles of habitat and competition and to forecast the future of the resident species. In addition, while these studies accounted for the imperfect detection of the focal species, no multi-season analysis of these species has accounted for the imperfect detection of the secondary species, potentially biasing inference. We analyzed survey data using models that combine the general multistate-multi-season occupancy modeling framework with autologistic modeling, allowing us to account for important aspects of our study system. We found that local extinction probability increases for each species when the other is present; however, the effect of the invader on the resident is greater. Although the species prefer different habitats, these habitats are highly correlated at the patch scale, and the impacts of invader on the resident are greatest in patches that would otherwise be optimal. As a consequence, competition leads to a weaker relationship between habitat and Northern Spotted Owl occupancy. Colonization and extinction rates of the invader are closely related to neighborhood occupancy, and over the first half of the study the availability of colonists limited the rate of population growth. Competition is likely to exclude the resident species, both through its immediate effects on local extinction and by indirectly lowering colonization rates as Northern Spotted Owl occupancy declines. Our analysis suggests that dispersal limitation affects both the invasion dynamics and the scale at which the effects of competition are observed. We also provide predictions regarding the potential costs and benefits of managing Barred Owl populations at different target levels.

Forest management guidelines for rare or declining species in the Pacific Northwest, USA, include both late successional reserves and specific vegetation management criteria. However, whether current management practices for well-studied species such as northern spotted owls (Strix occidentallis caurina) can aid in conserving a lesser known subspecies-Humboldt martens (Martes caurina humboldtensis)-is unclear. To address the lack of information for martens in coastal Oregon, USA, we quantified vegetation characteristics at locations used by Humboldt martens and spotted owls in two regions (central and southern coast) and at two spatial scales (the site level summarizing extensive vegetation surveys and regionally using remotely sensed vegetation and estimated habitat models). We estimated amount of predicted habitat for both species in established reserves. If predicted overlap in established reserves was low, then we reported vegetation characteristics to inform potential locations for reserves or management opportunities. In the Central Coast, very little overlap existed in vegetation characteristics between Humboldt martens and spotted owls at either the site or regional level. Humboldt martens occurred in young forests composed of small diameter trees with few snags or downed logs. Humboldt martens were also found in areas with very dense vegetation when overstory canopy and shrub cover percentages were combined. In the South Coast, Humboldt martens occurred in forests with smaller diameter trees than spotted owl sites on average. Coastal Humboldt martens may use stands of predicted high quality spotted owl habitat in the Pacific Northwest. Nonetheless, our observations suggest that coastal Humboldt martens exist in areas that include a much higher diversity of conifer size classes as long as extensive dense shrub cover, predominantly in the form of high salal and evergreen huckleberry, are available. We suggest that managers consider how structural characteristics (e.g., downed logs, shrub cover, patch size), are associated with long-term species persistence rather than relying on reserves based on broad cover types. Describing vegetation may partially describe suitability, but available prey or predation risk ultimately influence likelihood of individual Humboldt marten use. Guidelines for diversifying vegetation management, and retaining or restoring appropriate habitat conditions at both the stand level and regionally, may increase management flexibility and identify forest conditions that support both spotted owls and Humboldt martens.

Environmental regulations often require wildlife surveys prior to habitat disturbance to avoid impacts or as the basis for planning mitigation, yet project‐level surveys may not provide the insights needed to guide long‐term management. Management of the red tree vole (Arborimus longicaudus) has largely been based on such surveys. As an alternative approach, we evaluated distribution patterns using frequency of red tree vole occurrence and habitat suitability models to guide conservation planning. We developed a suite of models based on subsets of covariates from two previously developed models and evaluated the extent to which spatial covariates improved the models. We used presence–absence data that were collected from 364 randomly selected 1‐ha Current Vegetation Survey and Forest Inventory and Analysis plots to develop models and describe occurrence patterns. The best models included a spatial covariate, maximum tree diameter, distance from suitable habitat, forest age class, and the interaction between maximum tree diameter and forest age class. We compared performance of the previously published models, our best model, and an ensemble model that used predictions from all three models. Under the ensemble model, correct classification rates were relatively high and considerably improved, suggesting that the application of all three models provided greater accuracy than any individual model. We argue that habitat models, coupled with spatial patterns of the frequency of occurrence, can provide useful tools for addressing species management and may provide more insight than project‐level surveys. The use of habitat suitability models can therefore be closely tied to red tree vole management decisions and conservation strategies, as well as reducing survey costs that otherwise often make projects infeasible.

We describe the historical and current distribution of tree voles (Arborimus longicaudus; A. pomo) and compare the minimum density of trees with tree vole nests in different forest age-classes based on museum records, field notes of previous collectors, tree vole nest surveys conducted by federal agencies, and our field studies in Oregon and California. We conclude that tree voles are still fairly common in old forests within much of their historical range, but have become uncommon or rare in some areas as a result of fire and logging. Our analysis of food stored at red tree vole (A. longicaudus) nests in Oregon indicated that the vast majority of tree voles feed almost exclusively on needles of Douglas-fir (Pseudotsuga menziesii). However, tree voles in the Sitka spruce zone of northwestern Oregon feed primarily on western hemlock (Tsuga heterophylla) and Sitka spruce (Picea sitchensis). Historical field notes from the California coastal region also documented occasional voles that feed on grand fir (Abies grandis), Monterey pine (Pinus radiata), or Bishop pine (P. muricata). We used the program MaxEnt to develop predictive models of tree vole presence based on an a priori set of habitat, structure, and climate variables. Based on a comparison of potential vole habitat in historical and recent vegetation maps, we estimated that the geographic distribution of red tree voles in Oregon contracted by 23 percent in the period 1914–2013.

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.

Many membrane-bound proteins, including cytokines, receptors, and growth factors, are proteolytically cleaved to release a soluble form of their extracellular domain. The tumor necrosis factor (TNF)-alpha converting enzyme (TACE/ADAM-17) is a transmembrane metalloproteinase responsible for the proteolytic release or "shedding" of several cell-surface proteins, including TNF and p75 TNFR. We established a TACE-reconstitution system using TACE-deficient cells co-transfected with TACE and substrate cDNAs to study TACE function and regulation. Using the TACE-reconstitution system, we identified two additional substrates of TACE, interleukin (IL)-1R-II and p55 TNFR. Using truncations and chimeric constructs of TACE and another ADAM family member, ADAM-10, we studied the function of the different domains of TACE in three shedding activities. We found that TACE must be expressed with its membrane-anchoring domain for phorbol ester-stimulated shedding of TNF, p75 TNFR, and IL-1R-II, but that the cytoplasmic domain is not required for the shedding of these substrates. The catalytic domain of ADAM-10 could not be functionally substituted for that of TACE. IL-1R-II shedding required the cysteine-rich domain of TACE as well as the catalytic domain, whereas TNF and p75 TNFR shedding required only the tethered TACE catalytic domain.

Tumor necrosis factor-α (TNFα) is a cytokine that induces protective inflammatory reactions and kills tumor cells but also causes severe damage when produced in excess, as in rheumatoid arthritis and septic shock. Soluble TNFα is released from its membrane-bound precursor by a membrane-anchored proteinase, recently identified as a multidomain metalloproteinase called TNFα-converting enzyme or TACE. We have cocrystallized the catalytic domain of TACE with a hydroxamic acid inhibitor and have solved its 2.0 Å crystal structure. This structure reveals a polypeptide fold and a catalytic zinc environment resembling that of the snake venom metalloproteinases, identifying TACE as a member of the adamalysin/ADAM family. However, a number of large insertion loops generate unique surface features. The pro-TNFα cleavage site fits to the active site of TACE but seems also to be determined by its position relative to the base of the compact trimeric TNFα cone. The active-site cleft of TACE shares properties with the matrix metalloproteinases but exhibits unique features such as a deep S3′ pocket merging with the S1′ specificity pocket below the surface. The structure thus opens a different approach toward the design of specific synthetic TACE inhibitors, which could act as effective therapeutic agents in vivo to modulate TNFα-induced pathophysiological effects, and might also help to control related shedding processes.

The dihydrofolate reductase-deficient Chinese hamster ovary cell line, DXB11-CHO, commonly used as a host cell for the production of recombinant proteins requires 7.5% serum-supplementation for optimal growth. Regulatory issues surrounding the use of serum in clinical production processes and the direct and indirect costs of using serum in large-scale production and recovery processes have triggered efforts to derive serum-independent host cell lines. We have successfully isolated a serum-free host that we named Veggie- CHO. Veggie-CHO was generated by adapting DXB11-CHO cells to growth in serum-free media in the absence of exogenous growth factors such as Transferrin and Insulin-like growth factor, which we have previously shown to be essential for growth and viability of DXB11- CHO cells. Veggie-CHO cells have been shown to maintain an average doubling time of 22 hr in continuous growth cultures over a period of three months and have retained the dihydrofolate reductase -deficient phenotype of their parental DXB11-CHO cells. These properties and the stability of its serum-free phenotype have allowed the use of Veggie- CHO as host cells for transfection and amplified expression of recombinant proteins. We describe the derivation a serum-free recombinant cell line with an average doubling time of 20 hr and specific productivity of 2.5 Units recombinant Flt-3L protein per 10e6 cells per day.