Site Occupancy Dynamics of Northern Spotted Owls in the Eastern Cascades,Washington, USA, 1990–2003

Abstract: Northern spotted owls (Strix occidentalis caurina) have received intense research and management interest since their listing as a threatened species by the United States Fish and Wildlife Service in 1990. Several spotted owl (Strix occidentalis) response variables have been examined in various investigations, but recent advances in statistical modeling permit evaluations of temporal and spatial variability in site occupancy, local-extinction, and colonization probabilities while incorporating imperfect detection probabilities. Following recent work by other researchers on site occupancy dynamics of spotted owls in Oregon, USA, we evaluated temporal variability of detection, occupancy, local-extinction, and colonization probabilities for spotted owls, as well as potential influences of barred owl (Strix varia) presence on these parameters. We used spotted owl survey data collected from 1990 to 2003 on a study area in the eastern Cascades Mountains, Washington, USA, to compare competing occupancy models from Program PRESENCE using Akaike's Information Criterion. Detection probabilities for individual spotted owls ranged from 0.54 to 0.80 if barred owls were not detected during the survey season and from 0.19 to 0.71 if barred owls were detected during the survey season. Pair detection probabilities ranged from 0.27 to 0.67 if barred owls were not detected during an individual survey and from 0.09 to 0.36 if barred owls were detected during an individual survey. During the study, site occupancy probabilities for spotted owl pairs declined by approximately 50%. For all spotted owls, both singles and pairs, site occupancy probabilities declined moderately during the study. Barred owl presence was negatively associated with spotted owl detection probabilities, and it had a positive association with local-extinction probabilities for all spotted owls, both singles and pairs. Given that our study area has supported higher densities of barred owls for longer periods than other study areas, our results may provide insight into how barred owls have influenced spotted owl site occupancy dynamics in adjacent British Columbia, Canada, or will influence spotted owl site occupancy dynamics in Oregon and California, USA, in the future.     

Review of the effects of barred owls on spotted owls

Barred owls (Strix varia) are forest-dwelling owls, native to eastern North America, with populations that expanded westward into the range of the spotted owl (Strix occidentalis). Barred owls exert an overwhelmingly negative influence on spotted owls, thereby threatening spotted owl population viability where the species co-occur. In this review, we provide an overview of the barred owl's range expansion and detail and synthesize previously published literature on spotted and barred owls within the range of the spotted owl as related to potential future outcomes for the northern spotted owl (S. o. caurina). We include research on diet, habitat use and selection, effects of barred owls on spotted owl demography and behavior, hybridization with spotted owls, parasites, contemporary management, and future research needs for spotted owl populations given continued barred owl expansion throughout western North America. Our literature review and synthesis should provide managers with the information necessary to develop strategies that mitigate deleterious effects of barred owls at local and landscape scales. © 2019 The Wildlife Society.     

Using Detection Dogs to Conduct Simultaneous Surveys of Northern Spotted (Strix occidentalis caurina) and Barred Owls (Strix varia)

State and federal actions to conserve northern spotted owl (Strix occidentalis caurina) habitat are largely initiated by establishing habitat occupancy. Northern spotted owl occupancy is typically assessed by eliciting their response to simulated conspecific vocalizations. However, proximity of barred owls (Strix varia)-a significant threat to northern spotted owls-can suppress northern spotted owl responsiveness to vocalization surveys and hence their probability of detection. We developed a survey method to simultaneously detect both species that does not require vocalization. Detection dogs (Canis familiaris) located owl pellets accumulated under roost sites, within search areas selected using habitat association maps. We compared success of detection dog surveys to vocalization surveys slightly modified from the U.S. Fish and Wildlife Service's Draft 2010 Survey Protocol. Seventeen 2 km ×2 km polygons were each surveyed multiple times in an area where northern spotted owls were known to nest prior to 1997 and barred owl density was thought to be low. Mitochondrial DNA was used to confirm species from pellets detected by dogs. Spotted owl and barred owl detection probabilities were significantly higher for dog than vocalization surveys. For spotted owls, this difference increased with number of site visits. Cumulative detection probabilities of northern spotted owls were 29% after session 1, 62% after session 2, and 87% after session 3 for dog surveys, compared to 25% after session 1, increasing to 59% by session 6 for vocalization surveys. Mean detection probability for barred owls was 20.1% for dog surveys and 7.3% for vocal surveys. Results suggest that detection dog surveys can complement vocalization surveys by providing a reliable method for establishing occupancy of both northern spotted and barred owl without requiring owl vocalization. This helps meet objectives of Recovery Actions 24 and 25 of the Revised Recovery Plan for the Northern Spotted Owl.     

Detection and vocalisations of three owl species (Strigiformes) in temperate rainforests of southern Chile

Conspecific broadcasts are effective to increase detection of owls. To determine the most appropriate time of the year to survey owls, we played conspecific owl vocalisations monthly in a temperate rainforest of southern Chile. From 12 broadcast points surveyed we recorded detections of Glaucidium nana, Strix rufipes and Tyto alba. Glaucidium nana presented a bimodal detection curve throughout the year and we recorded two regular vocalisations in response to broadcasting: contact pair call and territorial call. Strix rufipes and T. alba both showed a peak of detection between February and May. Strix rufipes presented three vocalisations: territorial call, contact pair call and female contact pair call while T. alba uttered two vocalisations: territorial call and twittering call. We recommend surveys during the end of the breeding season (austral summer–autumn) when detection is higher in most owls. Surveys should also take into consideration the variability of the vocalisations and include covariates in monitoring to evaluate occupancy/detection models.     

Site occupancy dynamics of northern spotted owls in managed interior Douglas fir forests,California, USA, 1995–2009

Northern spotted owls (Strix occidentalis caurina) have received intense research and management interest since their listing as a threatened species by the United States Fish and Wildlife Service in 1990. For example, public and private forest managers in the Pacific Northwest, USA, conduct surveys to determine presence or absence of spotted owls prior to timber harvest operations. However, although recently developed statistical methods have been applied to presence–absence data collected during research surveys, the effectiveness of operational surveys for detecting spotted owls and evaluating site occupancy dynamics is not known. We used spotted owl survey data collected from 1995 to 2009 on a study area in interior northern California, USA, to evaluate competing occupancy models from Program PRESENCE using Akaike's Information Criterion (AIC). During 1,282 individual surveys, we recorded 480 spotted owl detections (37.4%) and 13 barred owl (1.0%) detections. Average per visit detection probability (85% CL) for single and paired spotted owls was 0.93 (0.90–0.96) for informed daytime, stand-based searches and 0.47 (0.43–0.51) for nighttime, station-based surveys (estimated from the best model); the average per visit detection probability from the null model was 0.67 (0.64–0.70). Average pair-only detection probabilities were 0.86 (0.81–0.90) for informed daytime, stand-based searches and 0.23 (0.18–0.29) for nighttime, station-based surveys; the average per visit detection probability from the null model was 0.63 (0.58–0.68). Site occupancy for any owl declined from 0.81 (0.59–0.93) in 1995 to 0.50 (0.39–0.60) in 2009; pair occupancy declined from 0.75 (0.56–0.87) to 0.46 (0.31–0.61). Our results suggest that a combination of 1 informed stand and 2 station-based operational surveys can support determinations of spotted owl site status (either a single or a pair) at desired levels of confidence. However, our information was collected in an area where barred owls were rarely detected. Surveys conducted in areas that support well-established barred owl populations are likely to be less effective for determining presence or absence of spotted owls and may require more surveys and/or different survey methods to determine site status with confidence. © 2012 The Wildlife Society.     

Barred Owl Space Use and Habitat Selection in the Eastern Cascades,Washington

ABSTRACT Competition with barred owls (Strix varia varia) is an important factor contributing to the continued decline of threatened northern spotted owl (Strix occidentalis caurina) populations in the Pacific Northwest, USA, but basic information on habitat selection and space use patterns of barred owls is lacking for much of the region. We investigated space use and habitat selection by tracking radiotagged barred owls in the Eastern Cascade Range of Washington, USA, from 2004 to 2006. We surveyed for barred owls across the 309-km² study area and confirmed presence of barred owl pairs at 21 sites. We collected movement data on 14 barred owls from 12 sites. Mean annual 95% fixed-kernel home-range size was 194 ha for females (n = 4, SD = 70) and 288 ha for males (n = 5, SD = 114). Home ranges were located more frequently than expected in areas with low topographic position, gentle slopes, large overstory tree-crown diameter, high normalized difference vegetation index (NDVI), overstory tree canopy closure >72%, and a moderate amount of solar insolation. Within home ranges, areas that had large tree-crown diameters, low topographic positions, and gentle slopes were used more frequently than expected. The resource selection function we developed for barred owls in our study area indicated that barred owls used areas with the combination of low values for topographic position and slope and higher values for NDVI, solar insolation, and an interaction term for canopy closure and tree-crown diameter. In comparison to published information on northern spotted owls, barred owls used areas with similar canopy closure and tree size classes, but barred owl home ranges were much smaller and more concentrated on gentler slopes in valley bottoms. This information may contribute to the development of management practices that maintain forest characteristics appropriate for spotted owl habitat and prey in areas where spotted owls are least likely to be excluded by territorial barred owls in the Eastern Cascades of Washington.     

Using bioacoustics to enhance the efficiency of spotted owl surveys and facilitate forest restoration

The California spotted owl (Strix occidentalis occidentalis) is an older-forest associated species that resides at the center of forest management planning in the Sierra Nevada and Southern California, USA, which are experiencing increasingly large and severe wildfires and drought-related tree mortality. We leveraged advances in passive acoustic survey technologies to develop an acoustically assisted survey design that could increase the efficiency and effectiveness of project-level surveys for spotted owls, allowing surveys to be completed in a single year instead of in multiple years. We deployed an array of autonomous recording units (ARUs) across a landscape and identified spotted owl vocalizations in the resulting audio using BirdNET. We then evaluated spatio-temporal patterns in spotted owl vocalizations near occupied territories and the ability of a crew naïve to the location of occupied territories to locate spotted owls based on patterns of acoustic detections. After only 3 weeks of acoustic surveys, ≥1 ARU within 750 m of all 17 occupied territories obtained spotted owl detections across ≥2 nights. When active surveys using broadcast calling were conducted near ARUs with spotted owl detections by surveyors naïve to territory occupancy status and locations, surveyors located owls in 93% to 100% of occupied territories with ≤3 surveys. To further improve the efficiency of spotted owl surveys, we developed a statistical model to identify and prioritize areas across the Sierra Nevada for different survey methods (active only, acoustically assisted, no surveys) based on the expected probability of occupancy predicted from remotely sensed measurements of tree height and historical occupancy. Depending on managers' tolerance for false negatives, this model could help identify large areas that might not benefit from surveys based on low expected occupancy probabilities and areas where acoustically assisted surveys might enhance survey effectiveness and efficiency. Collectively, these findings can help managers streamline the survey process and thus increase the pace of forest restoration while minimizing potential near-term adverse effects on California spotted owls.     

The Invasion of Barred Owls and its Potential Effect on the Spotted Owl: a Conservation Conundrum

The spotted owl (Strix occidentalis) is a threatened species in many areas of its western North American range. Concomitant with its decline has been a rapid invasion of its range and habitat by barred owls (Strix varia), a native species that was restricted, until relatively recently, to eastern North America. We assess the theoretical potential for negative interactions between these two owls by examining size dimorphism and ecological relationships within various owl assemblages throughout the world. We then review the anecdotal, natural history, modeling, and experimental evidence that suggest barred owls may negatively affect spotted owls with at least a potential for the competitive exclusion of spotted owls by barred owls throughout all or part of the former’2019;s range. While it is widely accepted that barred owls are either causing or exacerbating declines of spotted owl populations, there are confounding factors, such as habitat loss and bad weather that also may contribute to declines of spotted owls. Both theory and empirical information suggest that barred owls are likely to have negative effects on spotted owl range and density, but the degree of the impact is not predictable. There is a conservation conundrum here, in that the barred owl is a native species that has expanded its range westwards, either naturally or with a degree of human facilitation, and now constitutes a major threat to the viability of another native species, the threatened spotted owl. We propose that only through carefully designed experiments involving removal of barred owls will we be able to determine if recent declines in spotted owl populations are caused by barred owls or by other factors. It is rare in conservation science that replicate study areas exist for which we also have long-standing demographic information, as is the case with the spotted owl. Removal experiments would take advantage of the wealth of data on spotted owls, and allow ecologists to assess formally the impacts of an invasive species on a threatened species, as well as to suggest mitigation measures.     

Genetic structure, introgression, and a narrow hybrid zone between northern and California spotted owls (Strix occidentalis)

The northern spotted owl (Strix occidentalis caurina) is a threatened subspecies and the California spotted owl (Strix occidentalis occidentalis) is a subspecies of special concern in the western United States. Concern for their continued viability has arisen because of habitat loss caused by timber harvesting. The taxonomic status of the northern subspecies has been the subject of continuing controversy. We investigated the phylogeographical and population genetic structure of northern and California spotted owls with special reference to their region of contact. Mitochondrial DNA (mtDNA) control region sequences confirmed the existence of two well-differentiated lineages connected by a narrow hybrid zone in a region of low population density in north central California. Maximum-likelihood estimates indicated bidirectional gene flow between the lineages but limited introgression outside the region of contact. The lengths of both the mtDNA hybrid zone and the reduced density patch were similar and slightly exceeded estimates of natal dispersal distances. This suggests that the two subspecies were in secondary contact in a hybrid zone trapped by a population density trough. Consequently, the zone of interaction is expected to be geographically stable. We discovered a third, rare clade of haplotypes, which we interpreted to be a result of incomplete lineage sorting; those haplotypes result in a paraphyletic northern spotted owl with respect to the California spotted owl. A congeneric species, the barred owl (Strix varia), occasionally hybridizes with spotted owls; our results indicated an upper bound for the frequency of barred owl mtDNA haplotypes in northern spotted owl populations of 3%.     

Development of 37 microsatellite loci for the great gray owl (<Emphasis Type="Italic">Strix nebulosa</Emphasis>) and other <Emphasis Type="Italic">Strix</Emphasis> spp. owls

We developed 37 great gray owl (Strix nebulosa) microsatellite primers from CA and TAGA enriched genomic libraries. Primers were tested in 15 great gray owls from California, USA and Alberta, Canada as well as two other Strix species, spotted owl (S. occidentalis) and barred owl (S. varia). These markers will have broad application in investigations of Strix population structure and genetic diversity.     

Microsatellite loci for distinguishing spotted owls (Strix occidentalis), barred owls (Strix varia), and their hybrids

We identified four diagnostic microsatellite loci that distinguish spotted owls (Strix occidentalis), barred owls (Strix varia), F₁ hybrids and backcrosses. Thirty‐four out of 52 loci tested (65.4%) successfully amplified, and four of these loci (11.8%) had allele sizes that did not overlap between spotted and barred owls. The probability of correctly identifying a backcross with these four loci is 0.875. Genotyping potential hybrid owls with these markers revealed that field identifications were often wrong. Given the difficulty of identifying hybrids in the field, these markers will be useful for hybrid identification, law enforcement and spotted owl conservation.     

Relationship between wildfire,salvage logging,and occupancy of nesting territories by northern spotted owls

The northern spotted owl (Strix occidentalis caurina) is one of the most intensively studied raptors in the world; however, little is known about the impacts of wildfire on the subspecies and how they use recently burned areas. Three large-scale wildfires in southwest Oregon provided an opportunity to investigate the short-term impacts of wildfire and salvage logging on site occupancy of spotted owls. We used Program MARK to develop single-species, multiple-season models of site occupancy using data collected during demographic surveys of spotted owl territories. In our first analysis, we compared occupancy dynamics of spotted owl nesting territories before (1992–2002) and after the Timbered Rock burn (2003–2006) to a reference area in the south Cascade Mountains that was not affected recently by wildfire. We found that the South Cascades had greater colonization probabilities than Timbered Rock before and after wildfire ( , 95% CI = 0.60–2.03), and colonization probabilities declined over time at both areas ( , 95% CI = −0.12 to 0.00). Extinction probabilities were greater at South Cascades than at Timbered Rock prior to the burn ( , 95% CI = 0.23–2.62); however, Timbered Rock had greater extinction probabilities following wildfire ( , 95% CI = 0.29–2.62). The Timbered Rock and South Cascades study areas had similar patterns in site occupancy prior to the Timbered Rock burn (1992–2001). Furthermore, Timbered Rock had a 64% reduction in site occupancy following wildfire (2003–2006) in contrast to a 25% reduction in site occupancy at South Cascades during the same time period. This suggested that the combined effects of habitat disturbances due to wildfire and subsequent salvage logging on private lands negatively affected site occupancy by spotted owls. In our second analysis, we investigated the relationship between wildfire, salvage logging, and occupancy of spotted owl territories at the Biscuit, Quartz, and Timbered Rock burns from 2003 to 2006. Extinction probabilities increased as the combined area of early seral forests, high severity burn, and salvage logging increased within the core nesting areas ( , 95% CI = 0.10–3.66). We were unable to identify any relationships between initial occupancy or colonization probabilities and the habitat covariates that we considered in our analysis where the β coefficient did not overlap zero. We concluded that site occupancy of spotted owl nesting territories declined in the short-term following wildfire, and habitat modification and loss due to past timber harvest, high severity fire, and salvage logging jointly contributed to declines in site occupancy. © 2013 The Wildlife Society.     

Multistate Models Reveal Long-Term Trends of Northern Spotted Owls in the Absence of a Novel Competitor

Quantifying spatial and temporal variability in population trends is a critical aspect of successful management of imperiled species. We evaluated territory occupancy dynamics of northern spotted owls (Strix occidentalis caurina), California, USA, 1990–2014. The study area possessed two unique aspects. First, timber management has occurred for over 100 years, resulting in dramatically different forest successional and structural conditions compared to other areas. Second, the barred owl (Strix varia), an exotic congener known to exert significant negative effects on spotted owls, has not colonized the study area. We used a Bayesian dynamic multistate model to evaluate if territory occupancy of reproductive spotted owls has declined as in other study areas. The state-space approach for dynamic multistate modeling imputes the number of territories for each nesting state and allows for the estimation of longer-term trends in occupied or reproductive territories from longitudinal studies. The multistate approach accounts for different detection probabilities by nesting state (to account for either inherent differences in detection or for the use of different survey methods for different occupancy states) and reduces bias in state assignment. Estimated linear trends in the number of reproductive territories suggested an average loss of approximately one half territory per year (-0.55, 90% CRI: -0.76, -0.33), in one management block and a loss of 0.15 per year (-0.15, 90% CRI: -0.24, -0.07), in another management block during the 25 year observation period. Estimated trends in the third management block were also negative, but substantial uncertainty existed in the estimate (-0.09, 90% CRI: -0.35, 0.17). Our results indicate that the number of territories occupied by northern spotted owl pairs remained relatively constant over a 25 year period (-0.07, 90% CRI: -0.20, 0.05; -0.01, 90% CRI: -0.19, 0.16; -0.16, 90% CRI: -0.40, 0.06). However, we cannot exclude small-to-moderate declines or increases in paired territory numbers due to uncertainty in our estimates. Collectively, we conclude spotted owl pair populations on this landscape managed for commercial timber production appear to be more stable and do not show sharp year-over-year declines seen in both managed and unmanaged landscapes with substantial barred owl colonization and persistence. Continued monitoring of reproductive territories can determine whether recent declines continue or whether trends reverse as they have on four previous occasions. Experimental investigations to evaluate changes to spotted owl occupancy dynamics when barred owl populations are reduced or removed entirely can confirm the generality of this conclusion.     

Reproduction of northern spotted owls: The role of local weather and regional climate

We examined associations between annual reproduction and climate for 6 populations of individually marked northern spotted owls (Strix occidentalis caurina) in Washington and Oregon. We used an information-theoretical approach and mixed models to evaluate statistical models representing a priori hypotheses about the effects of weather and climate on reproduction. Reproduction was higher for adult than subadult owls and declined as the proportion of spotted owl territories with barred owl (Strix varia) detections increased. Similar to other spotted owl studies, we found that reproduction was negatively associated with cold, wet winters and nesting seasons at 3 of 6 study areas. In addition, we identified new relationships between reproduction, annual precipitation, storms, and regional climate cycles. For 3 of 6 areas, we found a quadratic relation between precipitation (rain and snow) and reproduction, with the number of young fledged per pair per year declining as precipitation in the previous year deviated from average levels. A meta-analysis conducted across all 6 areas indicated that reproduction at the regional level had a quadratic association with total winter snowfall in the preceding winter and was positively related to temperatures during the previous summer and fall. The amount of annual variation in reproduction accounted for by weather and climate varied widely across the 6 areas (4–79%), whereas variation in weather and climate across owl territories accounted for little of the spatial variation in reproduction (0–4%). Our results suggest that across the range of the species climate factors affecting prey abundance may have a greater effect on reproduction than direct effects of weather on nestlings. © 2011 The Wildlife Society.     

Subspecific relationships and genetic structure in the spotted owl

Hierarchical genetic structure was examined in the three geographically-defined subspecies of spotted owl (Strix occidentalis) to define relationships among subspecies and quantify variation within and among regional and local populations. Sequences (522 bp) from domains I and II of the mitochondrial control region were analyzed for 213 individuals from 30 local breeding areas. Results confirmed significant differences between northern spotted owls and the other traditional geographically defined subspecies but did not provide support for subspecific level differences between California and Mexican spotted owls. Divergence times among subspecies estimated with a 936 bp portion of the cytochrome b gene dated Northern and California/Mexican spotted owl divergence time to 115,000–125,000 years ago, whereas California/Mexican spotted owl divergence was estimated at 15,000 years ago. Nested clade analyses indicated an association between California spotted owl and Mexican spotted owl haplotypes, implying historical contact between the two groups. Results also identified a number of individuals geographically classified as northern spotted owls (S. o. caurina) that contained haplotypes identified as California spotted owls (S. o. caurina). Among all northern spotted owls sampled (n=131), 12.9% contained California spotted owl haplotypes. In the Klamath region, which is the contact zone between the two subspecies, 20.3% (n=59) of owls were classified as California spotted owls. The Klamath region is a zone of hybridization and speciation for many other taxa as well. Analyses of population structure indicated gene flow among regions within geographically defined subspecies although there was significant differentiation among northern and southern regions of Mexican spotted owls. Among all areas examined, genetic diversity was not significantly reduced except in California spotted owls where the southern region consists of one haplotype. Our results indicate a stable contact zone between northern and California spotted owls, maintaining distinct subspecific haplotypes within their traditional ranges. This supports recovery efforts based on the traditional subspecies designation for the northern spotted owl. Further, although little variation was found between California and Mexican spotted owls, we suggest they should be managed separately because of current isolation between groups.     

Effects of survey methods on burrowing owl behaviors

Monitoring wildlife populations often involves intensive survey efforts to attain reliable estimates of population size. Such efforts can increase disturbance to animals, alter detection, and bias population estimates. Burrowing owls (Athene cunicularia) are declining across western North America, and information on the relative effects of potential survey methods on owl behaviors is needed. We designed a field experiment to compare burrowing owl flight distances, times displaced, and probabilities of being displaced between 4 potential population survey methods (single walking surveyor, single vehicle stop, single vehicle stop with 2 surveyors, and double vehicle stop with 2 surveyors), and an experimental control in the agricultural matrix of Imperial Valley, California. Between 25 April and 1 May 2008, we randomly applied survey methods to 395 adult male owls during daylight hours (0700 hours through 1900 hours). All survey methods increased odds of displacing owls from perches. Survey methods with observers outside the vehicle were 3 times more likely to displace an owl than a single vehicle stop where observers remained inside the vehicle. Owls were displaced farther distances by all survey methods compared to control trials, but distances and time displaced did not differ among survey methods. We recommend that surveys for counting owls during the breeding season in agroecystems like the Imperial Valley where high densities of owls nest primarily along the borders of fields be conducted using single vehicle stops with or without 2 surveyors, depending on conditions for locating owls from roads. © 2011 The Wildlife Society.     

Empirical support for a despotic distribution in a California spotted owl population

Territorial species, such as the spotted owl (Strix occidentalis),are predicted to follow an ideal despotic distribution. However,debate exists on whether wild populations actually meet theassumptions of an ideal distribution, such as perfect perceptualabilities (i.e., the ability to recognize high- and low-qualitysites without error). Because this hypothesis has importantlife history ramifications for spotted owls, we investigatedwhether occupancy rates of California spotted owl (S. o. occidentalis)territories in the San Bernardino Mountains of southern Californiapositively correlated with a qualitative "potential fitness"(denoted by _pf) estimated from survival and reproduction ofterritorial owls. Spotted owls in our study tended to occupyterritories with the highest _pf, supporting the assumption ofideal perceptual abilities within this population. However,this relationship was noisy, and we suggest that some individualsdo not assess site quality accurately because of perceptuallimitations, prey dynamics, and large territory sizes. Furthermore,dispersal processes, high survival rates, and long life spansof spotted owls may be other key factors preventing some individualsfrom selecting sites of the highest quality and, consequently,our ability to precisely estimate _pf.     

Habitat selection by northern spotted owls in mixed-coniferous forests

Conservation planning for the federally threatened northern spotted owl (Strix occidentalis caurina) requires an ability to predict their responses to existing and future habitat conditions. To inform such planning we modeled habitat selection by northern spotted owls based upon fine-scale (approx. 1.0 ha) characteristics within stands comprised primarily of mixed-aged, mixed coniferous forests of southwestern Oregon and north-central California. We sampled nocturnal (i.e., primarily foraging) habitat use by 71 radio-tagged spotted owls over 5 yr in 3 study areas and sampled vegetative and physical environmental conditions at inventory plots within 95% utilization distributions of each bird. We compared conditions at available forest patches, represented by the inventory plots, with those at patches used by owls using discrete-choice regressions, the coefficients from which were used to construct exponential resource selection functions (RSFs) for each study area and for all 3 areas combined. Cross-validation testing indicated that the combined RSF was reasonably robust to local variation in habitat availability. The relative probability that a fine-scale patch was selected decreased nonlinearly with distances from nests and streams; varied unimodally with increasing average diameter of coniferous trees and also with increasing basal area of Douglas-fir (Pseudotsuga menziesii) trees; increased linearly with increasing basal areas of sugar pine (Pinus lambertiana) and hardwood trees and with increasing density of understory shrubs. Large-diameter trees (>66 cm) appeared important <400 m from nest sites. The RSF can support comparative risk assessments of the short- versus long-term effects of silvicultural alternatives designed to integrate forest ecosystem restoration and habitat improvement for northern spotted owls. Results suggest fine-scale factors may influence population fitness among spotted owls. © 2011 The Wildlife Society.     

Population decline in California spotted owls near their southern range boundary

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 ( 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.