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Breeding dispersal, the movement from one breeding territory to another, is rare for philopatric species that evolved within relatively stable environments, such as the oldgrowth coniferous forests of the Pacific Northwest. Although dispersal is not inherently maladaptive, the consequences of increased dispersal on population dynamics in populations whose historical dispersal rates are low could be significant, particularly for a declining species. We examined rates and possible causes of breeding dispersal based on a sample of 4,118 northern spotted owls (Strix occidentalis caurina) monitored in seven study areas over 28 yr, 1990–2017, in Oregon and Washington, USA. Using a multistate mark–resight analysis, we investigated the potential impacts of an emergent congeneric competitor (barred owl Strix varia) and forest alteration (extrinsic factors), and social and individual conditions (intrinsic factors) on 408 successive and 1,372 nonsuccessive dispersal events between years. The annual probability of breeding dispersal increased for individual owls that had also dispersed in the previous year and decreased for owls on territories with historically high levels of reproduction. Intrinsic factors including pair status, prior reproductive success, and experience at a site, were also associated with breeding dispersal movements. The percent of monitored owls dispersing each year increased from ∼˜7% early in the study to ∼25% at the end of the study, which coincided with a rapid increase in numbers of invasive and competitively dominant barred owls. We suggest that the results presented here can inform spotted owl conservation efforts as we identify factors contributing to changing rates of demographic parameters including site fidelity and breeding dispersal. Our study further shows that increasing rates of breeding dispersal associated with population declines contribute to population instability and vulnerability of northern spotted owls to extinction, and the prognosis is unlikely to change unless active management interventions are undertaken.

Invasive species are a primary threat to biodiversity and are challenging to manage once populations become established in previously unoccupied areas. But removing them is further complicated when invasions occur in continental, mixed-ownership systems. We demonstrate a rare conservation success: the regional-scale removal of an invasive predator — the barred owl (Strix varia) — to benefit the spotted owl (Strix occidentalis) in California. Barred owl site occupancy declined sixfold, from 0.19 to 0.03, following 1 year of removals, and site extinction (0.92) far exceeded colonization (0.02). Spotted owls recolonized 56% of formerly occupied territories within 1 year, contrasting starkly with removals conducted after barred owls achieved high densities in the Pacific Northwest. Our study therefore averted the otherwise likely extirpation of California spotted owls (Strix occidentalis occidentalis) by barred owl competition. Collectively, leveraging technological advances in population monitoring, early intervention, targeting defensible biogeographic areas, and fostering public—private partnerships will reduce invasive species-driven extinction of native fauna in continental systems.

Recent extensions of occupancy modeling have focused not only on the distribution of species over space, but also on additional state variables (e.g., reproducing or not, with or without disease organisms, relative abundance categories) that provide extra information about occupied sites. These biologist-driven extensions are characterized by ambiguity in both species presence and correct state classification, caused by imperfect detection. We first show the relationships between independently published approaches to the modeling of multistate occupancy. We then extend the pattern-based modeling to the case of sampling over multiple seasons or years in order to estimate state transition probabilities associated with system dynamics. The methodology and its potential for addressing relevant ecological questions are demonstrated using both maximum likelihood (occupancy and successful reproduction dynamics of California Spotted Owl) and Markov chain Monte Carlo estimation approaches (changes in relative abundance of green frogs in Maryland). Just as multistate capture—recapture modeling has revolutionized the study of individual marked animals, we believe that multistate occupancy modeling will dramatically increase our ability to address interesting questions about ecological processes underlying population-level dynamics.

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 (φ), and a reparameterization of the Jolly-Seber capture–recapture model (i.e. reverse Jolly-Seber; RJS) to estimate annual rates of population change (λRJS) 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 (λ) suggested that Spotted Owl populations declined from 1.2% to 8.4% per year depending on the study area. The weighted mean estimate of λ 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°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 Niño 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.

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).

Context Fire-adapted species have evolved to exploit resources in heterogenous landscapes that presumably maximize energy acquisition and minimize energetic expenditure. However, limited empirical work exists demonstrating the explicit energetic mechanisms that drive such adaptive responses to fire across diverse landscapes. Objectives The California spotted owl ( Strix occidentalis occidentalis ) appears to benefit from landscape heterogeneity and preferentially uses smaller patches of severely burned forest, a behavior that has been hypothesized as adaptive. Here, we investigate empirical support for this hypothesis. Methods We leveraged high-resolution GPS tracking and nest video monitoring to examine the hunting success, movement, and nest provisioning of 34 spotted owls in the Sierra Nevada and San Bernardino Mountains, California across burned and unburned landscapes. Results Regardless of time since fire, individuals avoided foraging directly within moderately or severely burned patches. 1 to 2 years post-fire, individuals had more success capturing prey in unburned forest, and the energy individuals spent moving increased with the proportion of high-severity fire and decreased with the proportion unburned forest. Multiple years after a fire, individuals had more success capturing prey, spent less energy moving, and provisioned more energy to nests in landscapes with more low-severity fire. Conclusions These results support the hypothesis that spotted owls are adapted to fire-prone landscapes and that disturbance events within this region’s natural range of variation can ultimately promote hunting and provisioning. As fires deviate from regional norms across the globe, the negative impacts of fire may become more extreme and long-term benefits of fire may degrade for animals in fire-prone landscapes. Examining the mechanistic impacts of disturbance can allow us to better understand animal responses to rapidly changing landscapes.

More frequent and extreme heat waves threaten climate-sensitive species. Structurally complex, older forests can buffer these effects by creating cool microclimates, although the mechanisms by which forest refugia mitigate physiological responses to heat exposure and subsequent population-level consequences remain relatively unexplored. We leveraged fine-scale movement data, doubly labeled water, and two decades of demographic data for the California spotted owl (Strix occidentalis occidentalis) to (1) assess the role of older forest characteristics as potential energetic buffers for individuals and (2) examine the subsequent value of older forests as refugia for a core population in the Sierra Nevada and a periphery population in the San Bernardino Mountains. Individuals spent less energy moving during warmer sampling periods and the presence of tall canopies facilitated energetic conservation during daytime roosting activities. In the core population, where tall-canopied forest was prevalent, temperature anomalies did not affect territory occupancy dynamics as warmer sites were both less likely to go extinct and less likely to become colonized, suggesting a trade-off between foraging opportunities and temperature exposure. In the peripheral population, sites were more likely to become unoccupied following warm summers, presumably because of less prevalent older forest conditions. While individuals avoided elevated energetic expenditure associated with temperature exposure, behavioral strategies to conserve energy may have diverted time and energy from reproduction or territory defense. Conserving older forests, which are threatened due to fire and drought, may benefit individuals from energetic consequences of exposure to stressful thermal conditions.

Management of many North American forests is challenged by the need to balance the potentially competing objectives of reducing risks posed by high-severity wildfires and protecting threatened species. In the Sierra Nevada, California, concern about high-severity fires has increased in recent decades but uncertainty exists over the effects of fuel-reduction treatments on species associated with older forests, such as the California Spotted Owl ( Strix occidentalis occidentalis ). Here, we assessed the effects of forest conditions, fuel reductions, and wildfire on a declining population of Spotted Owls in the central Sierra Nevada using 20 years of demographic data collected at 74 Spotted Owl territories. Adult survival and territory colonization probabilities were relatively high, while territory extinction probability was relatively low, especially in territories that had relatively large amounts of high canopy cover (≥70%) forest. Reproduction was negatively associated with the area of medium-intensity timber harvests characteristic of proposed fuel treatments. Our results also suggested that the amount of edge between older forests and shrub/sapling vegetation and increased habitat heterogeneity may positively influence demographic rates of Spotted Owls. Finally, high-severity fire negatively influenced the probability of territory colonization. Despite correlations between owl demographic rates and several habitat variables, life stage simulation (sensitivity) analyses indicated that the amount of forest with high canopy cover was the primary driver of population growth and equilibrium occupancy at the scale of individual territories. Greater than 90% of medium-intensity harvests converted high-canopy-cover forests into lower-canopy-cover vegetation classes, suggesting that landscape-scale fuel treatments in such stands could have short-term negative impacts on populations of California Spotted Owls. Moreover, high-canopy-cover forests declined by an average of 7.4% across territories during our study, suggesting that habitat loss could have contributed to declines in abundance and territory occupancy. We recommend that managers consider the existing amount and spatial distribution of high-canopy forest before implementing fuel treatments within an owl territory, and that treatments be accompanied by a rigorous monitoring program.

Vocal territory defense can vary within a species due to many factors such as sex and breeding status, influencing territory size and thus population density across a landscape. Therefore, understanding what influences variation in territorial vocalizations can help to illuminate trade‐offs between territoriality and other life history demands, which benefits our general understanding of animal ecology as well as helps to inform emerging passive acoustic monitoring approaches. Here, we investigated how sex and breeding status affected territoriality and vocal behavior in the California spotted owl Strix occidentalis occidentalis in the Sierra Nevada, California, USA, using high‐resolution acoustic/GPS tags. We discovered that territorial vocal behavior was related to breeding status and to a lesser extent sex. Breeding owls with fledged young had a less diverse vocal repertoire, produced fewer and quieter territorial calls, and typically called only when close to their nest. Males were also more likely to engage in territorial calling than females. Breeding spotted owls also maintained significantly smaller territories – but utilized larger home ranges – than non‐breeding individuals. Our results suggest that breeding spotted owls may reduce their investment in territorial behaviors to mitigate the demands and risks associated with rearing young. Further, our results have important implications for the passive acoustic monitoring of spotted owls and, more broadly, highlight the utility of using multiple call types to detect species of interest.

Species worldwide have begun to shift their range boundaries in response to climate change and other anthropogenic causes, with population declines at the trailing edge of a species’ range often foreshadowing future changes in core parts of the range. Therefore, we analyzed a 30-year (1991–2019) data set for the California spotted owl (Strix occidentalis occidentalis) near its southern range boundary in southern California, USA, that included the largest regional population (San Bernardino Mountains) to estimate trends in territory occupancy and reproduction. We then assessed how these demographic rates were affected by habitat, wildfire, fuel treatments, and climate. Mean occupancy declined from 0.82 to 0.39 during our study, whereas reproductive output showed no temporal trends (x̄ = 0.65 young/occupied territory). Territory extinction (extirpation) rates were relatively low in territories with more large trees (≥50 cm dbh), and colonization increased strongly with large tree density for low-elevation territories within the shrub-woodland ecotype but not for higher-elevation territories within mixed-conifer forest. High-severity wildfire had an adverse effect on occupancy: territory extinction rates steadily increased with the amount of high-severity fire within an owl territory during the previous 10 years, while colonization declined to nearly zero when ≥40% of a territory burned at high-severity during the previous 10 years. The effects of high-severity fire were unlikely to be confounded with post-fire fuel treatments, which primarily consisted of the removal, burning, or scattering of brush and small trees and snags (<40.6 cm dbh) and affected much smaller areas than high-severity fire. Of the 40 territories that received fuel treatments within 10 years of a fire, only 3 of them had post-fire fuel treatments that affected >5% of the territory, whereas average area burned at high severity for all 40 territories was 17%. Fuel treatments intended to modify fire behavior and reduce the likelihood of large, high-severity fires led to increases in territory extinction and colonization such that their net effect on occupancy was minimal. Our simulations of occupancy dynamics indicated that high-severity fire accounted for 9.6% of the observed decline in occupancy, whereas fuel treatments effectively accounted for none of the decline. Spotted owl reproductive output was lower at territories where fuel treatments occurred, but low- to moderate-severity fire resulted in much larger, population-level reductions in reproductive output (141 fewer young) from 2006–2019 than treatments (19 fewer young). Thus, the benefits of fuel treatments that reduce fire occurrence and severity appear to outweigh potential short-term costs to spotted owls and their habitat. Because high-severity fire only explained a modest amount of the long-term occupancy decline and much of the decline occurred in the 1990s before large fires occurred, additional factors are likely adversely affecting the owl population and merit further study. Nevertheless, the large observed population decline, limited evidence of owl dispersal among mountain ranges in the southern California metapopulation, and negative effects of increasingly large and severe fire suggest that California spotted owls at their southern range boundary are vulnerable to extirpation. In an era of climate change, owls in the core part of the range will likely become increasingly susceptible to warmer temperatures and increased severe fire activity in the future. Thus, the restoration of historical, low-severity fire regimes through fuels management while maintaining large trees is important to improving owl persistence.