A preliminary investigation of the spotted owl in Oregon

Between 1970 and 1974, data were collected on the distribution and biology of the spotted owl (Strix occidentalis) in Oregon. One-hundred and sixteen pairs and seven single birds were located. Spotted owls occurred throughout the mountains of western Oregon and on the east slope of the Cascade Range at least as far east as Badger Butte, Hood River County; Abbot Butte, Jefferson County; and Swan Lake Point, Klamath County. The upper elevational limits of the species increased from about 1,350 meters in northern Oregon to 1,770 meters in southern Oregon. Although spotted owls were not uncommon in some areas, evidence indicated that the population was declining as a result of habitat loss. A total of 2,647 prey items were identified from 42 pairs of owls. Prey species included 29 mammals, 20 birds, 2 reptiles, a crayfish, a terrestrial snail, and 26 genera of insects. Mammalian prey comprised over 90 percent of the biomass consumed. The flying squirrel (Glaucomys sabrinus) was the principal prey species (13-48 percent of total biomass consumed), except in dry forest areas, where wood rats (Neotoma fuscipes and N. cinerea) became most important (7-78 percent of total biomass). Other important prey included snowshoe hares (Lepus americanus), red tree voles (Phenacomys longicaudus), deer mice (Peromyscus maniculatus), western red-backed voles (Clethrionomys occidentalis), Mazama pocket gophers (Thomomys mazama), pikas (Ochotona princeps), and small birds. Predation on showshoe hares, gophers, moles, and insects was heaviest during the late spring and summer months. Spotted owls foraged primarily at night, and often captured arboreal animals (squirrels, wood rats, and tree voles) by grabbing them from limbs or tree trunks. Eighteen spotted owl nests were located, including 13 in cavities in living old-growth conifers, three in clumps of deformed limbs caused by dwarfmistletoes (Arceuthobium spp. ), and two in platform nests constructed by other species. Nest height above ground ranged from 19. 8-55.2 meters. Owls added no materials to their nests except small amounts of molted down. The mean date of clutch initiation for 15 nests was 29 March (range 9 March - 19 April). As egg-laying neared, adult activities (vocalization, copulation, courtship feeding, roosting) became increasingly centered around nest trees, and several days before they laid eggs, females began to roost inside their nest cavities. Incubation, which lasted approximately 30 days, was performed entirely by females. Males fed females during this period. Owlets fledged at 34-36 days of age (between early May and mid -June), and were fed by their parents until late September. At fledging, owlets were weak fliers, and often fell from their nests to the ground. When this occurred, they regained safe perches in trees or low bushes by climbing. Forty-eight nesting attempts were observed (38 successful). Total number of young fledged was 63. Mean number of young fledged per successful nest was 1.61 (range 1-3). The percentage of pairs attempting to nest ranged from 89 percent in 1972, to 16 percent in 1973, and 46 percent in 1974. I suspected, but did not verify, that the low numbers of breeding pairs in some years reflected a decline in prey numbers. Percent of nesting pairs which fledged young was 92 in 1972, 40 in 1973, and 72 in 1974. A principal cause of nest failure in all years was nest desertion during early stages of nesting. In 1972, juvenile mortality between fledging and the end of August was 35 percent. Predation was suspected as the principal cause of mortality, but several young were killed when the fell from nests, and two young died in their nest. Most broods did not move far from their nests until they began dispersal in September or October. Of 14 broods checked in late August, 5 were within 160 meters of their nests, 5 were 170-250 meters from their nests, and 2 had moved 487 and 670 meters, respectively. Two broods could not be relocated, but had moved 1, 050 and 365 meters, respectively, when last seen in July. Owlets underwent two molts during their first summer; the white natal down was replaced by the downy mesoptile plumage before owlets fledged. Replacement of the mesoptile plumage by the first winter plumage then occurred over a 4-month period and was complete by the end of September or early October. In the latter plumage owlets were nearly indistinguishable from adults. Of 123 sites where spotted owls were located, 117 (95 percent) where characterized by unharvested old-growth conifer forests. Two pairs occupied old-growth forests which had been partially logged about 30 years previous, and three occupied second-growth forests which contained minor old -growth components. The multilayered structure of old-growth stands provided owls with large trees for nests and winter roosts, small shaded summer roost trees, and a closed canopy (canopy closure ranged from 53-86 percent at nest sites). Owls occurred in most coniferous associations found in western Oregon and the Cascades, with the exception of subalpine forests, open ponderosa pine (Pinus ponderosa) forests and lodgepole pine (Pinus contorta) forests. Owls showed a slight preference (X2 = P < . 01) for nests located on north or east exposures, possibly because trees there were usually larger and forests were denser than on south or west exposures. Fifteen of 18 nests were within 400 meters of permanent water (range = 15-1,417 meters). Sources of water utilized by all 18 pairs consisted of small perennial streams or springs. Timber harvest occurred or was scheduled in 52 percent of the owl habitats located during the study. In most cases, timber harvest within an occupied habitat did not drive the owls completely out of the area, because only small portions of extensive forest areas were harvested. When portions of small forest areas (less than about 80 hectares) were harvested, however, owls often disappeared from these areas. Two pairs located in old-growth forests which had been subjected to very light over story removal indicated that, under some circumstances, owls could tolerate this type of harvest activity. Clear-cut harvest, however, eliminated roosting, nesting and most foraging in the affected areas

Population trends in northern spotted owls: Associations with climate in the Pacific Northwest

We used reverse time capture-mark-recapture models to describe associations between rate of population change (λ) and climate for northern spotted owls (Strix occidentalis caurina) at six long-term study areas in Washington and Oregon, USA. Populations in three of six areas showed strong evidence of declining populations, while populations in two additional areas were likely declining as well. At four areas, λ was positively associated with wetter-than-normal conditions during the growing season, which likely affects prey availability. Lambda was also negatively associated with cold, wet winters and nesting seasons, and the number of hot summer days. The amount of annual variation in λ accounted for by climate varied across study areas (3–85%). Rate of population change was more sensitive to adult survival than to recruitment; however, there was considerable variation among years and across study areas for all demographic rates. While annual survival was more closely related to regional climate conditions, recruitment was often associated with local weather. In addition to climate, declines in recruitment at four of six areas were associated with increased presence of barred owls. Climate change models predict warmer, wetter winters and hotter, drier summers for the Pacific Northwest in the first half of the 21st century. Our results indicate that these conditions have the potential to negatively affect annual survival, recruitment, and consequently population growth rates for northern spotted owls

Terpenoid Resin Distribution in Conifer Needles with Implications for Red Tree Vole, Arborimus longicaudus, Foraging

Tree voles are dietary specialists, feeding almost exclusively on conifer needles and bark. They reduce their exposure to conifer chemical defenses by physically removing resin ducts from many needles before ingesting the remaining tissue. The portion of needle removed differs among tree species, depending on the location of the resin ducts. To evaluate the amount of resin avoided by this behavior we removed the resin ducts from Douglas-fir, Western Hemlock, and Sitka Spruce needles and used gas chromatography to compare volatile resin concentrations in needles with and without the resin ducts removed. Needle tissues without resin ducts contained no terpenoid resin, demonstrating that tree voles can regulate the nutritional quality of their diet by controlling the amount of resin ingested. We suggest that differences in the physical structure (and possibly chemical composition of terpenes) of the needles make it difficult for voles to easily switch between tree hosts

Population trends in northern spotted owls: Associations with climate in the Pacific Northwest

We used reverse time capture-mark-recapture models to describe associations between rate of population change (λ) and climate for northern spotted owls (Strix occidentalis caurina) at six long-term study areas in Washington and Oregon, USA. Populations in three of six areas showed strong evidence of declining populations, while populations in two additional areas were likely declining as well. At four areas, λ was positively associated with wetter-than-normal conditions during the growing season, which likely affects prey availability. Lambda was also negatively associated with cold, wet winters and nesting seasons, and the number of hot summer days. The amount of annual variation in λ accounted for by climate varied across study areas (3–85%). Rate of population change was more sensitive to adult survival than to recruitment; however, there was considerable variation among years and across study areas for all demographic rates. While annual survival was more closely related to regional climate conditions, recruitment was often associated with local weather. In addition to climate, declines in recruitment at four of six areas were associated with increased presence of barred owls. Climate change models predict warmer, wetter winters and hotter, drier summers for the Pacific Northwest in the first half of the 21st century. Our results indicate that these conditions have the potential to negatively affect annual survival, recruitment, and consequently population growth rates for northern spotted owls.This is the author's final (post-refereeing) manuscript. Article appears in Biological Conservation (www.elsevier.com/ locate/biocon) and is copyrighted by Elsevier

Competitive Interactions and Resource Partitioning Between Northern Spotted Owls and Barred Owls in Western Oregon

The federally threatened northern spotted owl (Strix occidentalis caurina) is the focus of intensive conservation efforts that have led to much forested land being reserved as habitat for the owl and associated wildlife species throughout the Pacific Northwest of the United States. Recently, however, a relatively new threat to spotted owls has emerged in the form of an invasive competitor: the congeneric barred owl (S. varia). As barred owls have rapidly expanded their populations into the entire range of the northern spotted owl, mounting evidence indicates that they are displacing, hybridizing with, and even killing spotted owls. The range expansion by barred owls into western North America has made an already complex conservation issue even more contentious, and a lack of information on the ecological relationships between the 2 species has hampered recovery efforts for northern spotted owls. We investigated spatial relationships, habitat use, diets, survival, and reproduction of sympatric spotted owls and barred owls in western Oregon, USA, during 2007–2009. Our overall objective was to determine the potential for and possible consequences of competition for space, habitat, and food between these previously allopatric owl species. Our study included 29 spotted owls and 28 barred owls that were radio-marked in 36 neighboring territories and monitored over a 24-month period. Based on repeated surveys of both species, the number of territories occupied by pairs of barred owls in the 745-km² study area (82) greatly outnumbered those occupied by pairs of spotted owls (15). Estimates of mean size of home ranges and core-use areas of spotted owls (1,843 ha and 305 ha, respectively) were 2–4 times larger than those of barred owls (581 ha and 188 ha, respectively). Individual spotted and barred owls in adjacent territories often had overlapping home ranges, but interspecific space sharing was largely restricted to broader foraging areas in the home range with minimal spatial overlap among core-use areas. We used an information-theoretic approach to rank discrete-choice models representing alternative hypotheses about the influence of forest conditions, topography, and interspecific interactions on species-specific patterns of nighttime resource selection. Spotted owls spent a disproportionate amount of time foraging on steep slopes in ravines dominated by old (>120 yr) conifer trees. Barred owls used available forest types more evenly than spotted owls, and were most strongly associated with patches of large hardwood and conifer trees that occupied relatively flat areas along streams. Spotted and barred owls differed in the relative use of old conifer forest (greater for spotted owls) and slope conditions (steeper slopes for spotted owls), but we found no evidence that the 2 species differed in their use of young, mature, and riparian-hardwood forest types. Mean overlap in proportional use of different forest types between individual spotted owls and barred owls in adjacent territories was 81% (range=30–99%). The best model of habitat use for spotted owls indicated that the relative probability of a location being used was substantially reduced if the location was within or in close proximity to a core-use area of a barred owl. We used pellet analysis and measures of food-niche overlap to determine the potential for dietary competition between spatially associated pairs of spotted owls and barred owls. We identified 1,223 prey items from 15 territories occupied by spotted owls and 4,299 prey items from 24 territories occupied by barred owls. Diets of both species were dominated by nocturnal mammals, but diets of barred owls included many terrestrial, aquatic, and diurnal prey species that were rare or absent in diets of spotted owls. Northern flying squirrels (Glaucomys sabrinus), woodrats (Neotoma fuscipes, N. cinerea), and lagomorphs (Lepus americanus, Sylvilagus bachmani) were primary prey for both owl species, accounting for 81% and 49% of total dietary biomass for spotted owls and barred owls, respectively. Mean dietary overlap between pairs of spotted and barred owls in adjacent territories was moderate (42%; range=28–70%). Barred owls displayed demographic superiority over spotted owls; annual survival probability of spotted owls from known-fate analyses (0.81, SE=0.05) was lower than that of barred owls (0.92, SE=0.04), and pairs of barred owls produced an average of 4.4 times more young than pairs of spotted owls over a 3-year period. We found a strong, positive relationship between seasonal (6-month) survival probabilities of both species and the proportion of old (>120 yr) conifer forest within individual home ranges, which suggested that availability of old forest was a potential limiting factor in the competitive relationship between these 2 species. The annual number of young produced by spotted owls increased linearly with increasing distance from a territory center of a pair of barred owls, and all spotted owls that attempted to nest within 1.5 km of a nest used by barred owls failed to successfully produce young. We identified strong associations between the presence of barred owls and the behavior and fitness potential of spotted owls, as shown by changes in movements, habitat use, and reproductive output of spotted owls exposed to different levels of spatial overlap with territorial barred owls. When viewed collectively, our results support the hypothesis that interference competition with barred owls for territorial space can constrain the availability of critical resources required for successful recruitment and reproduction of spotted owls. Availability of old forests and associated prey species appeared to be the most strongly limiting factors in the competitive relationship between these species, indicating that further loss of these conditions can lead to increases in competitive pressure. Our findings have broad implications for the conservation of spotted owls, as they suggest that spatial heterogeneity in vital rates may not arise solely because of differences among territories in the quality or abundance of forest habitat, but also because of the spatial distribution of a newly established competitor. Experimental removal of barred owls could be used to test this hypothesis and determine whether localized control of barred owl numbers is an ecologically practical and socio-politically acceptable management tool to consider in conservation strategies for spotted owls.Keywords: Home range, Strix occidentalis caurina, Strix varia, Resource partitioning, Competition, Survival, Barred owl, Niche overlap, Reproduction, Northern spotted ow

Roosting Habitat Use and Selection By Northern Spotted Owls During Natal Dispersal

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.Keywords: Dispersal habitat, Radio-telemetry, Washington, Logistic regression, Northern spotted owl, Strix occidentalis caurin

Home Range and Habitat Selection by Northern Spotted Owls on the Eastern Slope of the Cascade Mountains, Washington

We used radiotelemetry to study space use and habitat selection of 16 Northern Spotted Owls (Strix occidentalis caurina) on the eastern slope of the Cascade Mountains, Washington, U.S.A., in 1989–1990. We used a geographical information system (GIS) and aerial photo interpretation of digital orthophotos to assign owl locations a value for vegetation type, topographic position, amount of edge, and distance to water. We compared owl relocations and random locations within 95% fixed kernel (FK) home ranges to determine each owl's selection of cover types, using logistic regression and generalized estimating equations (GEE) to estimate an exponential resource selection function likelihood. Minimum convex polygon (MCP) home ranges (SE) averaged 2858 ha (712 ha) for males and 1883 ha (249 ha) for females. Individual 95% FK home ranges averaged 1980 ha (229 ha) for males and 1649 ha (163 ha) for females. Pair home ranges averaged 3419 ha (826 ha) for MCP and 2427 ha (243 ha) for 95% FK. Nonbreeding season home ranges averaged approximately 3.5 times larger than breeding season home ranges for both males and females. Our best habitat model indicated that owls selected closed-canopy forests with a component of large (≥50 cm dbh) trees for roosting and foraging. In a given cover type, owls foraged lower on the slope. Management circles centered on nest areas—commonly used as a surrogate for home ranges—can be relatively poor representations of actual ranges used by pairs. However, an alternative for managing Spotted Owl home ranges is not readily available. Maintaining sufficient closed-canopy forest to provide habitat for Spotted Owls in the dry, fire-prone forests on the eastern slope of the Washington Cascades will be a challenge because forestry methods used to reduce the risk or severity of fire generally reduce the prevalence of structural features that characterize good Spotted Owl habitat.Keywords: Northern Spotted Owl, fixed kernel, Washington, minimum convex polygon, radiotelemetry, Strix occidentalis caurina, habitat selection, home rang

Variation in inbreeding rates across the range of Northern Spotted Owls (\u3ci\u3eStrix occidentalis caurina\u3c/i\u3e): Insights from over 30 years of monitoring data

Inbreeding has been difficult to quantify in wild populations because of incomplete parentage information. We applied and extended a recently developed framework for addressing this problem to infer inbreeding rates in Northern Spotted Owls (Strix occidentalis caurina) across the Pacific Northwest, USA. Using pedigrees from 14,187 Northern Spotted Owls, we inferred inbreeding rates for 14 types of matings among relatives that produce pedigree inbreeding coefficients of F=0.25 or F=0.125. Inbreeding was most common in the Washington Cascades, where an estimated 15% of individuals are inbred. Inbreeding was lowest in western Oregon (3.5%) and northern California (2.7%), and intermediate for the Olympic Peninsula of Washington (6.1%). Estimates from the Olympic Peninsula were likely underestimates because of small sample sizes and the presence of few pedigrees capable of resolving inbreeding events. Most inbreeding resulted from matings between full siblings or half siblings, although a high rate of inbreeding from mother–son pairs was identified in the Olympic Peninsula. Geographic variation in inbreeding rates may reflect population declines and bottlenecks that have been detected in prior investigations. We show that there is strong selection against inbred birds. Only 3 of 44 inbred birds were later identified as parents (6.8%), whereas 2,823 of 10,380 birds that represented a comparable cross section of the data were later seen as reproducing parents (27.2%). Habitat loss and competition with Barred Owls (S. varia) remain primary threats to Northern Spotted Owls. However, given the negative consequences of inbreeding, Spotted Owl populations in Washington with suitable habitat and manageable numbers of Barred Owls may benefit from translocations of individuals from Oregon and California to introduce new genetic variation and reduce future inbreeding events. La endogamia ha sido dif´ıcil de cuantificar en las poblaciones silvestres debido a la falta de informaci ´on sobre los parentescos. Aplicamos y extendimos un marco conceptual recientemente desarrollado para encarar el problema de inferir las tasas de endogamia en Strix occidentalis caurina a trav´es del noroeste del Pac´ıfico, EEUU. Usando los pedigr´ıes provenientes de 14187 individuos, inferimos las tasas de endogamia para 14 tipos de apareamiento entre parientes que producen coeficientes de endogamia de pedigr´ı de F=0.25 o F=0.125. La endogamia fue ma´s com´un en las Cascadas de Washington, donde se estima que 15% de los individuos son endoga´micos. La endogamia fue menor en el oeste de Oreg´on (3.5%) y el norte de California (2.7%), e intermedia en la Pen´ınsula Ol´ımpica de Washington (6.1%). Las estimaciones de la Pen´ınsula Ol´ımpica fueron probablemente subestimadas debido a los peque ˜nos tama ˜nos de muestreo y a la presencia de pocos pedigr´ıes capaces de resolver los eventos de endogamia. La mayor´ıa de la endogamia result ´o de los apareamientos entre hermanos completos o medios hermanos, aunque se identific ´o una alta tasa de endogamia en parejas madre/hijo en la Pen´ınsula Ol´ımpica. La variaci ´on geogra´ fica en las tasas de endogamia puede reflejar disminuciones poblacionales y cuellos de botella que han sido detectados en investigaciones previas. Mostramos que hay una fuerte selecci ´on contra las aves endoga´micas. Solo tres de 44 aves endoga´micas fueron ma´s tarde identificadas como progenitores (6.8%), mientras que 2823 de 10380 aves que representaron una secci ´on transversal comparable de datos fueron vistas ma´s tarde como progenitores reproductivos (27.2%). La p´erdida de ha´bitat y la competencia con Strix varia sigue siendo la principal amenaza para S. o. caurina. Sin embargo, dadas las consecuencias negativas de la endogamia, las poblaciones de S. occidentalis en Washington con ha´bitat adecuado y n´umeros manejables de Strix varia pueden beneficiarse de traslocaciones de individuos de Oreg´on y California para introducir nueva variaci ´on gen´etica y reducir futuros eventos de endogamia

The past and future roles of competition and habitat in the range-wide occupancy dynamics of Northern Spotted Owls

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)

Range-wide sources of variation in reproductive rates of northern spotted owls

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