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.     

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%.     

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.     

Barred owl occupancy surveys within the range of the northern spotted owl

The range expansion by barred owls (Strix varia) into western North America has raised considerable concern regarding their potential effects on declining northern spotted owl (Strix occidentalis caurina) populations, yet most information on the occurrence of barred owls in the region is limited to incidental detections during surveys for spotted owls. To address this shortcoming we investigated response behavior, detection probabilities, and landscape occupancy patterns of barred owls in western Oregon, USA, during conspecific versus spotted owl call-broadcast surveys. Subtle differences in barred owl response behavior to conspecific versus spotted owl vocalizations combined with minor procedural differences between species-specific survey protocols led to a sizeable difference in estimated detection probabilities during conspecific (0.66, 95% CI = 0.61–0.71) versus spotted owl (0.48, 95% CI = 0.39–0.56) surveys. We identified 61 territorial pairs of barred owls during repeated surveys of a multi-ownership study area with the probability of occupancy being highest in the structurally diverse mixture of mature and old forests that occurred almost entirely on public lands. Our findings suggest that research and management strategies to address potential competitive interactions between spotted owls and barred owls will require carefully designed, species-specific survey methods that account for erratic response behaviors and imperfect detection of both species. Our sampling methods can be used by forest managers to determine the occurrence and distribution of barred owls with high confidence. © 2011 The Wildlife Society.     

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.     

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.     

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.     

Evidence for recent population bottlenecks in northern spotted owls (<Emphasis Type="Italic">Strix occidentalis caurina</Emphasis>)

The northern spotted owl (Strix occidentalis caurina) is one of the most controversial threatened subspecies ever listed under the US Endangered Species Act. Despite protection of its remaining forest habitat, recent field studies show continued declines of northern spotted owls. One potential threat to northern spotted owls which has not yet been shown is loss of genetic variation from population bottlenecks. Bottlenecks can increase the probability of mating among related individuals, potentially causing inbreeding depression, and can decrease adaptive potential. Here we report evidence for recent bottlenecks in northern spotted owls using a large genetic dataset (352 individuals and 10 microsatellite loci). The signature of bottlenecks was strongest in the Washington Cascade Mountains, in agreement with field data. Our results provide independent evidence that northern spotted owls have recently declined, and suggest that loss of genetic variation is an emerging threat to the subspecies’ persistence. Reduced effective population size (N _e) shown here in addition to field evidence for demographic decline highlights the increasing vulnerability of this bird to extinction.     

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.     

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.     

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.     

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.     

Contrasting patterns of mitochondrial and microsatellite genetic structure among Western European populations of tawny owls (Strix aluco)

A recent study of mitochondrial phylogeography of tawny owls (Strix aluco) in western Europe suggested that this species survived the Pleistocene glaciations in three allopatric refugia located in Iberia, Italy, and the Balkans, and the latter was likely the predominant source of postglacial colonization of northern Europe. New data from seven microsatellite loci from 184 individual owls distributed among 14 populations were used to assess the genetic congruence between nuclear and mitochondrial DNA (mtDNA) markers. Microsatellites corroborated the major phylogeographical conclusions reached on the basis of the mtDNA sequences, but also showed important differences leading to novel inferences. Microsatellites corroborated the three major refugia and supported the Balkan origin of northern populations. When corrected for differences in effective population size, microsatellites and mtDNA yielded generally congruent overall estimates of population structure (N*ST=0.12 vs. RST=0.16); however, there was substantial heterogeneity in the RST among the seven nuclear loci that was not correlated with heterozygosity. Populations representing the Balkans postglacial expansion interact with populations from the other two refugia forming two clines near the Alps and the Pyrenees. In both cases, the apparent position of the contact zones differed substantially between markers due to the genetic composition of populations sampled in northern Italy and Madrid. Microsatellite data did not corroborate the lower genetic diversity of northern, recently populated regions as was found with mtDNA; this discrepancy was taken as evidence for a recent bottleneck recovery. Finally, this study suggests that congruence among genetic markers should be more likely in cases of range expansion into new areas than when populations interact across contact zones.     

Home range and habitat selection of spotted owls in the central Sierra Nevada

We studied home range and habitat selection of radio-marked adult California spotted owls (Strix occidentalis occidentalis) randomly selected from among the breeding population of owls in the central Sierra Nevada, California from June to October 2006. The most parsimonious home-range estimate for our data was 555 ha (SE = 100 ha). Home-range size was positively correlated with the number of vegetation patches in the home range (habitat heterogeneity). We used resource selection ratios to examine selection of vegetation types by owls within our study area. Owl home ranges contained a high proportion of mature conifer forest, relative to its availability, although the confidence interval for this estimate overlapped one. We also used resource selection functions (RSF) to examine owl foraging habitat selection. Relative probability of selection of foraging habitat was correlated with vegetation classes, patch size, and their interaction. Owls showed highest selection rates for large patches (>10 ha) of pole-sized coniferous forest. Our results suggested that spotted owls in the central Sierra Nevada used habitat that contained a high proportion of mature conifer forest at the home-range scale, but at a finer scale (foraging site selection) owls used other vegetation classes interspersed among mature forest patches, consistent with our hypothesis that spotted owls may use other forest types besides old growth and mature forests when foraging. Our study provides an unbiased estimate of habitat use by spotted owls in the central Sierra Nevada. Our results suggest that forest managers continue to protect remaining mature and old-growth forests in the central Sierra Nevada because owl home ranges contain high proportions of these habitats. However, our results also showed that owls used younger stands as foraging habitat so that landscape heterogeneity, with respect to cover types, may be an important consideration for management but we did not attempt to relate our findings to fitness of owls. Thus management for some level of landscape heterogeneity for the benefit of owls should proceed with caution or under an adaptive management framework. © 2011 The Wildlife Society.     

Microsatellite markers for the Mexican spotted owl (Strix occidentalis lucida)

A genomic cosmid library was used to develop seven highly polymorphic microsatellite markers for the Mexican spotted owl (Strix occidentalis lucida). These are the first reported microsatellite markers derived from this species. The cloned and sequenced repeat motifs include a triplet repeat of (AAT)_n, two tetranucleotide repeats of (GATA)_n, a tetranucleotide repeat of (ATCC)_n, a compound repeat of (GA)_n(GATA)_n and the two pentanucleotide repeats (AGAAT)_n and (ATTTT)_n. The microsatellites described represent six presumably independent loci with the two pentanucleotide repeats having originated from a single cosmid. Primer pairs allow locus‐specific amplification of each marker from Mexican spotted owl genomic DNA.     

Predictive modeling for allopatric Strix (Strigiformes: Strigidae) owls in South America: determinants of their distributions and ecological niche‐based processes

Strix (Strigidae) is a worldwide genus of 17 owl species typical of forested habitats, including Rusty‐barred Owls (S. hylophila), Chaco Owls (S. chacoensis), and Rufous‐legged Owls (S. rufipes) in South America. These species are distributed allopatrically, but the ecological traits that determine their distributions remain largely unknown and their phylogenetic relationships are unclear. We used species distribution models (SDMs) to identify variables explaining their distribution patterns and test hypotheses about ecological divergence and conservatism based on niche overlap analysis. For Rusty‐barred Owls and Chaco Owls, climatic factors related to temperature played a major role, whereas a rainfall variable was more important for Rufous‐legged Owls. When niche overlaps were compared, accounting for regional similarities in the habitat available to each species, an ecological niche divergence process was supported for Chaco Owl‐Rusty‐barred Owl and Chaco Owl‐Rufous‐legged Owl, whereas a niche conservatism process was supported for Rusty‐barred Owl‐Rufous‐legged Owl. Different ecological requirements support current species delimitation, but they are in disagreement with the two main hypotheses currently envisaged about their phylogenetic relationships (Chaco Owls as the sister taxa of either Rufous‐legged Owls or Rusty‐barred Owls) and support a new phylogenetic hypothesis (Rufous‐legged Owls as sister taxa of Rusty‐barred Owls). Our findings suggest that speciation of Rusty‐barred Owls and Rufous‐legged Owls was a vicariant event resulting from Atlantic marine transgressions in southern South America in the Miocene, but their niche was conserved because habitat changed little in their respective ranges. In contrast, Chaco Owls diverged ecologically from the other two species as a result of their adaptations to the habitat they currently occupy. Ecological and historical approaches in biogeography can be embedded to explain distribution patterns, and results provided by SDMs can be used to infer historical and ecological processes in an integrative way.     

Does variation of sex ratio enhance reproductive success of offspring in tawny owls (Strix aluco)

Tawny owls, Strix aluco, laid female-biased clutches on territories with more abundant prey (field voles) in June, the month that chicks fledge. This appeared to enhance the subsequent reproductive success of fledglings, as in 1995 there was a significant correlation between the number of chicks fledged by adult females and the June vole abundance in the territory on which they were reared as chicks. This relationship did not hold for males. Since tawny owls lay eggs in March, these results indicate that owls are able to predict the June vole numbers on their territory, and respond by producing more of the sex most likely to gain a long-term benefit when resources are good.     

Analysis of genetic parentage in the tawny owl ( Strix aluco ) reveals extra-pair paternity is low

We have investigated genetic parentage in a Swiss population of tawny owls (Strix aluco). To this end, we performed genetic analysis for six polymorphic loci of 49 avian microsatellite loci tested for cross-species amplification. We found one extra-pair young out of 137 (0.7%) nestlings in 37 families (2.7%). There was no intra-specific brood parasitism. Our results are in accordance with previous findings for other raptors and owls that genetic monogamy is the rule. Female tawny owls cannot raise offspring without a substantial contribution by their mates. Hence one favoured hypothesis is that high paternal investment in reproduction selects for behaviour that prevents cuckoldry. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.