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Predator populations are imperiled globally, due in part to changing habitat and trophic interactions. Theoretical and laboratory studies suggest that heterogeneous landscapes containing prey refuges acting as source habitats can benefit both predator and prey populations, although the importance of heterogeneity in natural systems is uncertain. Here, we tested the hypothesis that landscape heterogeneity mediates predator–prey interactions between the California spotted owl (Strix occidentalis occidentalis)—a mature forest species—and one of its principal prey, the dusky‐footed woodrat (Neotoma fuscipes)—a younger forest species—to the benefit of both. We did so by combining estimates of woodrat density and survival from live trapping and very high frequency tracking with direct observations of prey deliveries to dependent young by owls in both heterogeneous and homogeneous home ranges. Woodrat abundance was ~2.5 times higher in owl home ranges (14.12 km2) featuring greater heterogeneity in vegetation types (1805.0 ± 50.2 SE) compared to those dominated by mature forest (727.3 ± 51.9 SE), in large part because of high densities in young forests appearing to act as sources promoting woodrat densities in nearby mature forests. Woodrat mortality rates were low across vegetation types and did not differ between heterogeneous and homogeneous home ranges, yet all observed predation by owls occurred within mature forests, suggesting young forests may act as woodrat refuges. Owls exhibited a type 1 functional response, consuming ~2.5 times more woodrats in heterogeneous (31.1/month ± 5.2 SE) versus homogeneous (12.7/month ± 3.7 SE) home ranges. While consumption of smaller‐bodied alternative prey partially compensated for lower woodrat consumption in homogeneous home ranges, owls nevertheless consumed 30% more biomass in heterogeneous home ranges—approximately equivalent to the energetic needs of producing one additional offspring. Thus, a mosaic of vegetation types including young forest patches increased woodrat abundance and availability that, in turn, provided energetic and potentially reproductive benefits to mature forest‐associated spotted owls. More broadly, our findings provide strong empirical evidence that heterogeneous landscapes containing prey refuges can benefit both predator and prey populations. As anthropogenic activities continue to homogenize landscapes globally, promoting heterogeneous systems with prey refuges may benefit imperiled predators. [ABSTRACT FROM AUTHOR]

Context: The concept of landscape heterogeneity is central to species conservation; yet understanding the processes by which heterogeneity affects species can be challenging in practice. Complex and sometimes difficult-to-measure responses of species may reflect the outcome of life-history trade-offs shaped by different landscape properties. Objectives: We tested the hypothesis that a mosaic of forest stand types affected hunting and breeding success for California spotted owls (Strix occidentailis occidentalis). Methods: We integrated high-temporal-resolution GPS tags, video monitoring of nest sites, long-term assessments of reproductive status, and high-resolution remotely sensed vegetation data in a mixed-ownership landscape in the Sierra Nevada, California to test our hypothesis. Results: Spotted owls made shorter nocturnal movements in homogenous territories with large areas of medium-aged forest apparently because this forest type allowed direct movement paths to foraging sites. However, spotted owls delivered prey at a higher rate to nest sites when they had more forest edge in their territory, which presumably provided greater access to large-bodied woodrat (Neotoma spp.) prey. Further, spotted owl reproductive output was relatively high in heterogenous territories containing a mix of mature and open forest. Conclusions: The benefits heterogenous forests provide to hunting success appeared to outweigh costs associated with additional commuting distance to foraging sites and provided potential fitness benefits to spotted owls. We suggest that the effects of landscape heterogeneity can vary not only among, but also within, species and can reflect the outcome of trade-offs among different life history activities. Understanding the effects of landscape properties on biological communities will benefit from additional empirical and mechanistic studies of individual species. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Snow insulates the soil from air temperature, decreasing winter cold stress and altering energy use for organisms that overwinter in the soil. As climate change alters snowpack and air temperatures, it is critical to account for the role of snow in modulating vulnerability to winter climate change. Along elevational gradients in snowy mountains, snow cover increases but air temperature decreases, and it is unknown how these opposing gradients impact performance and fitness of organisms overwintering in the soil. We developed experimentally validated ecophysiological models of cold and energy stress over the past decade for the montane leaf beetle Chrysomela aeneicollis, along five replicated elevational transects in the Sierra Nevada mountains in California. Cold stress peaks at mid‐elevations, while high elevations are buffered by persistent snow cover, even in dry years. While protective against cold, snow increases energy stress for overwintering beetles, particularly at low elevations, potentially leading to mortality or energetic tradeoffs. Declining snowpack will predominantly impact mid‐elevation populations by increasing cold exposure, while high elevation habitats may provide refugia as drier winters become more common. [ABSTRACT FROM AUTHOR]

Mountain ecosystems contain strong elevational gradients in climate and vegetation that shape species distributions and the structure of animal communities. Nevertheless, studies of habitat selection for individual species rarely account for such gradients that often result in species being managed uniformly across their range, which may not improve conservation as intended. Therefore, we characterized variation in nocturnal habitat selection by 18 GPS‐tagged California spotted owls (Strix occidentalis occidentalis) along a 1400‐m elevational gradient in the Sierra Nevada, California. We characterized three‐dimensional forest structure with light detection and ranging data that we used in mixed‐effects resource‐ and step‐selection analyses of owl habitat selection. At lower elevations, owls selected stands with shorter trees, sites closer to hard edges between tall forests and open areas, sites with less diversity in forest seral types and sites with more ridge and southwest aspects. In contrast, owls at higher elevations selected the opposite. Within public forests that had taller trees and within their home range core (45% kernel density estimate of GPS points) areas, owls selected forests with less and more canopy cover at low and high elevations, respectively. Outside of their core areas, owls selected areas with fewer and more tall trees at low and high elevations, respectively. These findings may be explained by elevational gradients in prey distribution and variation in owl diet because owls consume more woodrats (Neotoma spp; earlier seral species) at lower elevations and more flying squirrels (Glaucomys sabrinus; older forest species) at higher elevations. Thus, at low elevations and in areas unlikely to support nesting, spotted owls could benefit from management that promotes woodrat habitat by encouraging oak regeneration and creating small brushy openings within forests with shorter (younger) trees. Conversely, at higher elevations, (1) enhancing flying squirrel habitat by promoting large trees and denser canopy on mesic sites and (2) managing for greater cover type diversity on southwest‐facing slopes and ridgetops is more likely to improve foraging habitat quality for spotted owls. The patterns of owl selection over elevational gradients has not been explicitly considered in most habitat management plans but clearly would improve management throughout mountain ecosystems. [ABSTRACT FROM AUTHOR]

Coordination between nuclear and mitochondrial genomes is critical to metabolic processes underlying animals' ability to adapt to local environments, yet consequences of mitonuclear interactions have rarely been investigated in populations where individuals with divergent mitochondrial and nuclear genomes naturally interbreed. Genetic variation in the leaf beetle Chrysomela aeneicollis was assessed along a latitudinal thermal gradient in California's Sierra Nevada. Variation at mitochondrial cytochrome oxidase II (COII) and the nuclear gene phosphoglucose isomerase (PGI) shows concordance and was significantly greater along a 65 km transect than 10 other loci. STRUCTURE analyses using neutral loci identified a southern and northern subpopulation, which interbreed in the central drainage Bishop Creek. COII and PGI were used as indicators of mitochondrial and nuclear genetic variation in field and laboratory experiments conducted on beetles from this admixed population. Fecundity, larval development rate, running speed and male mating frequency were higher for beetles with geographically "matched" than "mismatched" mitonuclear genotypes. Effects of mitonuclear mismatch were largest for individuals with northern nuclear genotypes possessing southern mitochondria and were most pronounced after heat treatment or at high elevation. These findings suggest that mitonuclear incompatibility diminishes performance and reproductive success in nature, effects that could intensify at environmental extremes. [ABSTRACT FROM AUTHOR]

• We examined California spotted owl resource selection in relationship to forest fuel measurements and used these results along with the relative stand density index to predict probability of foraging use and fire risk across the Sierra Nevada. • Owls selected for higher canopy base height, lower ladder fuel densities, and lower canopy bulk densities, all of which are consistent with goals to decrease fire risk through fuels reduction treatments. • Owls also selected for higher surface fuels and higher canopy layer count. • Most of the Sierra Nevada landscape could be classified as either low priority (low owl use/high forest resilience) or fuels treatment priority (low owl use/low forest resilience), regardless of ownership type. • Focusing on the synergies between fuels reduction treatment goals and owl resource selection preferences will likely allow managers to simultaneously reduce fire risk and improve spotted owl foraging habitat. Wildfire has been an important force in shaping biological diversity in forests of western North America. Patterns of reburning helped to maintain heterogenous landscapes with low tree densities that naturally limited the extent of severe stand-replacing fires. However, anthropogenic impacts over the past two centuries have dramatically altered fire activity and its ecological function, creating conditions that promote large and intense fires with no historical analog. Fuels reduction treatments represent proven methods to mitigate extreme fire events, but there is controversy over their potential effects to sensitive wildlife species. The California spotted owl (Strix occidentalis occidentalis) is a forest species that evolved under a frequent-fire regime. Controversy over how spotted owls may be impacted by fuels reduction is one of numerous contributing factors that has limited the pace and scale of implementation. To examine the relationships between spotted owls and fuels reduction treatments, we studied breeding season nocturnal foraging habitat selection related to fuel structural metrics of 159 GPS-tagged California spotted owls across the Sierra Nevada bioregion. Spotted owls selected for higher canopy base height, lower ladder fuel density, and lower canopy bulk density, which represent synergies between fuels reduction treatments and owl space use. Spotted owls also selected for higher surface fuels and higher canopy layer counts, representing potential trade-offs between fuels reduction goals and spotted owl space use. We overlaid the probability of space use by foraging spotted owls with relative stand density index (SDI), a measure of forest resilience, to classify Sierra Nevada landscapes into four groups: low management priority (low owl use/high forest resilience), fuels reduction priority (low owl use/low forest resilience), potential conflict zones (high owl use/low forest resilience), and habitat retention priority (high owl use/high forest resilience). Fuels reduction priority (34.4%) was the largest category, suggesting that potential impacts of fuels reduction on spotted owl foraging habitat may be less widespread in the Sierra Nevada than previously believed. Our results suggest that fuels reduction treatments, especially those focused on mutually beneficial goals such as increasing canopy base height, reducing ladder fuels, and reducing canopy bulk density may directly improve spotted owl foraging habitat while also mitigating the effects of large and severe wildfires on forest species and forest ecosystems. [ABSTRACT FROM AUTHOR]

Highlights • We studied California spotted owl demography in mixed-ownership landscapes. • Forest conditions and topography better-explained variation in data than ownership. • Occupancy and reproduction were higher in study areas with more private lands. • Differences between study areas due to elevation and young forest with hardwoods. • Including private lands in conservation research provides valuable insights. Abstract Biodiversity conservation in mixed-ownership landscapes often depends on contributions from privately-owned lands, where natural resource development can alter and produce novel habitat conditions for species of conservation concern. A lack of research on private lands stemming from access issues and concerns over regulatory outcomes, however, often limits evaluation of the impact of land management. The California spotted owl (Strix occidentalis occidentalis), for example, often occurs in mixed-ownership landscapes but research on this species has occurred primarily on public lands. Therefore, we conducted the first large-scale private-public cooperative and comparative analyses of California spotted owls inhabiting mixed-ownership landscapes in the Sierra Nevada, California, USA. We surveyed 151 spotted owl sites from 2013 to 2017 within two study systems: one comprised primarily of public lands (national forests) where the owl population has declined over the last ∼20 years and a set of study areas comprised mostly of private lands on which relatively high estimates of site occupancy were recently reported. Multistate occupancy modeling indicated that the probability of occupancy and successful reproduction by owls depended on site status in the previous year, with both probabilities highest at sites where owls successfully reproduced in the previous year, intermediate at occupied sites where owls had not successfully reproduced, and lowest at previously unoccupied sites. Site occupancy probability was higher at low-elevation sites and lower at sites that contained more open area and younger forest. Successful reproduction by owls was also more likely at low-elevation sites and at sites with more north-facing slope and younger forest with high basal area of hardwoods. Study areas with more private lands tended to occur at lower elevations and have greater amounts of younger forest with high basal area of hardwoods, which may have contributed to higher occupancy and reproductive probabilities than the study area with more public land. Thus, differences in occupancy and reproductive probabilities between study areas appeared to be the result of differences in topographic and vegetation conditions that likely promote populations of key spotted owl prey species. Our results suggest that private lands in mixed-ownership landscapes may contribute to spotted owl conservation by conferring different benefits to owls than public lands and, more broadly, highlight the importance of including private lands in conservation research and planning. [ABSTRACT FROM AUTHOR]

Since the 1970s, California Spotted Owls ( Strix occidentalis occidentalis [CSO]) have been documented on private forest lands currently owned by Sierra Pacific Industries (SPI) in the Sierra Nevada of California. In 2012, SPI began an occupancy study on a portion of the CSO population known to occur on or near its managed forests in 5 watershed study areas averaging 110 km2 each. These watersheds occur from the southern end of the Cascade Range to just north of Yosemite National Park. We concluded that 57 historical CSO sites existed in these areas at the beginning of the study. During 2012 through 2016, the survey effort increased the total number of known CSO sites in the study areas to 65. During the same period, the yearly occupancy of the CSO sites within the study areas ranged from 70 to 86%, with 98% of the sites occupied during at least 1 y. Crude densities during the study period were calculated to be 0.117 CSO-occupied sites km−2. Compared to other studies in the Sierra Nevada, the CSO populations on our 5-yr study areas showed relatively high occupancy rates and a wide range of crude densities. [ABSTRACT FROM AUTHOR]

ABSTRACT The great gray owl ( Strix nebulosa) is listed by the state of California as endangered, with a population estimate of fewer than 300 individuals in the state. Nest-site availability has been suggested as a limiting factor for population growth in California, but information on nest types and nesting habitat has been based on a small number of nests that may not fully represent the variety of conditions used by the species. We collated all known nesting records in the Sierra Nevada mountains of California since 1973 ( n = 56) and then visited 47 of the nest sites to characterize habitat and compare them with paired reference sites. Great gray owls used a diversity of trees (8 species) and nest types. Although great gray owls in California are considered conifer-forest specialists, 30% of nests were in oak trees and 21% were below 1,000 m, which loosely corresponds to the lower conifer-zone limit. Across all elevations and tree species, degree of deterioration was the most important factor differentiating nest trees from paired reference trees at the same meadow, with nest trees being significantly more decayed. Nest trees (mean dbh = 100.5 ± SD 30.3 cm) were also significantly larger than reference trees. Canopy cover within 50 m of nest trees ( [ABSTRACT FROM AUTHOR]

Invasive predators can have detrimental impacts on native species and biological communities through direct consumptive effects and indirect effects on trophic interactions, yet these relations are often poorly understood through the initial stages of predator expansion. Leveraging early, lethal collection efforts to study invasive barred owls (Strix varia) at the leading edge of their range expansion in northeastern California, we conducted DNA metabarcoding of intestinal samples to assess the diet of this invasive predator. Through the development of customized primers and this novel approach to observing owl diet, we screened the intestinal contents of 124 barred owls and detected a broad diet of 78 unique prey types (48 vertebrates and 30 invertebrates), including prey types undetected in previous methodologies. Mammals were the most consumed vertebrate class (frequency of occurrence = 65 %), followed by amphibians (32 %), birds (22 %), and reptiles (19 %). Diets differed regionally but were similar among ages and sexes and exhibited limited variation in response to local environmental conditions. Our work highlights the generalist predatory strategy of invasive barred owls, identifies numerous native species potentially threatened by their range expansion, and indicates that they will not serve as ecological replacements for congeneric spotted owls (Strix occidentalis) they displace. These findings suggest that expanding currently limited barred owl removals could benefit native species and communities in western North America. More broadly, we demonstrate DNA metabarcoding, combined with early intervention, provides a powerful tool for conducting detailed assessments of species consumed by invasive predators, potentially incentivizing conservation actions and improving outcomes. • We developed custom primers to assess the diet of an invasive predator. • We identified 78 unique prey types at a leading edge of an invasion. • We demonstrate higher taxonomic resolution than traditional methods. • We show how early invasive intervention can aid scientific inquiry. • We show that continued barred owl removals may benefit native species. [ABSTRACT FROM AUTHOR]