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.
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 (
), on the population dynamics of an iconic old-forest native species, the northern spotted owl (
). 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.
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.
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).
The federally threatened northern spotted owl (Strix occidentalis caurina) has been intensively studied across its range, and habitat needs for the species have influenced forest management in ...northwestern North America for decades. Dense forest canopies are often reported in the scientific literature and agency management plans as an important habitat attribute for spotted owls, though the means of measuring forest canopy and interpreting species requirements vary across studies and more importantly, among management plans. We used light detection and ranging (lidar) measurements of canopy cover, canopy surface heterogeneity, and upper canopy surface connectivity, and an index of the presence of a competitive invasive species, the barred owl (S. varia), in multinomial discrete choice models using a Bayesian framework to evaluate selection of forest cover types by spotted owls in Oregon, USA, 2008–2015. We designated yearly activity centers based on the most biologically significant observation during the nesting season (Mar–Aug), generally centered on the nest tree. Spotted owls selected activity centers with more canopy cover and higher heterogeneity of the canopy surface within 100 m than was available within their territories. The average proportion of canopy cover within 100 m of a spotted owl activity center was 0.79 ± 0.12 (SD; range = 0.34–0.99). The presence of barred owls did not explain variability in selection of spotted owl activity centers, but barred owls might not affect third-order habitat selection within territories, or our index was too spatially coarse to detect these effects on spotted owl resource selection. We demonstrate that lidar provides researchers and managers with a tool that can accurately measure forest canopies over large areas, and assist in mapping spotted owl habitat.
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.
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.
The frequency, extent, and severity of wildfire strongly influence the structure and function of ecosystems. Mixed‐severity fire regimes are the most complex and least understood fire regimes, and ...variability of fire severity can occur at fine spatial and temporal scales, depending on previous disturbance history, topography, fuel continuity, vegetation type, and weather. During high fire weather in 2013, a complex of mixed‐severity wildfires burned across multiple ownerships within the Klamath‐Siskiyou ecoregion of southwestern Oregon where northern spotted owl (Strix occidentalis caurina) demographics were studied since 1990. A year prior to these wildfires, high‐resolution, remotely sensed forest structural information derived from light detection and ranging (lidar) data was acquired for an area that fully covered the extent of these fires. To quantify wildfire impact on northern spotted owl nesting/roosting habitat, we fit a relative habitat suitability model based on pre‐fire locations used for nesting and roosting, and forest structure variables developed from 2012 lidar data. Our pre‐fire habitat suitability model predicted nesting/roosting locations well, and variable response functions followed known resource selection patterns. These forests had typical characteristics of old‐growth forest, with high density of large live trees, high canopy cover, and complex structure in canopy height. We projected the pre‐fire model onto lidar data collected two months post‐fire to produce a post‐fire suitability map, which indicated that >93% of pre‐fire habitat that burned at high severity was no longer suitable forest for nesting and roosting. We also quantified the probability that pre‐fire nesting/roosting habitat would burn at each severity class (unburned/low, low, moderate, high). Pre‐fire nesting/roosting habitat had lower probability of burning at moderate or high severity compared to other forest types under high burning conditions. Our results indicate that northern spotted owl habitat can buffer the negative effects of climate change by enhancing biodiversity and resistance to high‐severity fires, which are predicted to increase in frequency and extent with climate change. Within this region, protecting large blocks of old forests could be an integral component of management plans that successfully maintain variability of forests in this mixed‐ownership and mixed‐severity fire regime landscape and enhance conservation of many species.
We studied habitat selection by northern spotted owls (Strix occidentalis caurina) during natal dispersal in Washington State, USA, at both the roost site and landscape scales. We used logistic ...regression to obtain parameters for an exponential resource selection function based on vegetation attributes in roost and random plots in 76 forest stands that were used for roosting. We used a similar analysis to evaluate selection of landscape habitat attributes based on 301 radio-telemetry relocations and random points within our study area. We found no evidence of within-stand selection for any of the variables examined, but 78% of roosts were in stands with at least some large (>50 cm dbh) trees. At the landscape scale, owls selected for stands with high canopy cover (>70%). Dispersing owls selected vegetation types that were more similar to habitat selected by adult owls than habitat that would result from following guidelines previously proposed to maintain dispersal habitat. Our analysis indicates that juvenile owls select stands for roosting that have greater canopy cover than is recommended in current agency guidelines. Published 2014. This article is a U.S. Government work and is in the public domain in the USA.
PDF
