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Scott A. Gremel

Publications and source records attributed to Scott A. Gremel.

8 recordsLinked to original sources

Listening for extinction: Range-wide occupancy highlights critical risks and priorities for northern spotted owls

The northern spotted owl ( Strix occidentalis caurina ) continues to decline across its range, threatened by habitat disturbances and invasive barred owls ( Strix varia ). We conducted a range-wide spotted owl occupancy assessment using 2.1 million h of passive acoustic recordings from 4081 survey stations within 1027 randomly selected 5-km 2 hexagons across six physiographic regions. Applying multistate occupancy models, we estimated probabilities of landscape use (by at least one northern spotted owl) and pair occupancy (occupied by both a male and a female), while accounting for imperfect and sex-biased detection probability. Detection probability of pairs was low in most regions, primarily due to undetected females, but increased with survey effort. If management decisions can only be made when there is a high probability of detecting both sexes when they are present, then longer surveys are one approach to support the decisions. Landscape use and pair occupancy were positively associated with old-growth forest structure and topographic diversity. We also found a pronounced latitudinal gradient in spotted owl landscape use rates with estimated mean landscape use of 0.17 (SE = 0.03) and 0.16 (SE = 0.02) in the two most northern regions, and 0.54 (SE = 0.04) and 0.52 (SE = 0.05) in the two most southern regions. Pair occupancy rates were higher in southern regions (0.29, SE = 0.03, 0.43, SE = 0.05) and very low in northern regions (0.04, SE = 0.01, 0.03, SE = 0.01). In contrast to spotted owls, barred owls were detected in 90% of northern hexagons and 50% of southern hexagons, and barred owl calling intensity was over eight times higher than spotted owls. These results highlight extinction risks in several regions and suggest that management actions, such as spatially targeted barred owl control and protection of structurally complex habitats, have a narrow temporal window in which they can be effective in lowering the probability of regional extirpations. We generated predictive maps of landscape use and pair occupancy, which could help guide regional conservation strategies. This study demonstrates how passive acoustic monitoring and machine learning can be used for broad-scale ecological monitoring, identifying extinction risk thresholds, supporting adaptive management, and improving conservation outcomes for wide-ranging and elusive species that can reliably be detected through unique vocalizations.

California, Oregon, Washington

Estimating northern spotted owl (Strix occidentalis caurina) pair detection probabilities based on call-back surveys associated with long-term mark-recapture studies, 1993–2018

The northern spotted owl ( Strix occidentalis caurina ; hereinafter NSO) was listed as “threatened” under the Endangered Species Act in 1990 and population declines have continued since that listing. Given the species’ protected status, any proposed activities on Federal lands that might impact NSO require consultation with U.S. Fish and Wildlife Service and part of that consultation often includes surveys to determine presence and occupancy status of the species in the proposed activity area. The objective of this report is to present study-area specific estimates of the probability of detection for NSO pairs from twelve 2-week seasonal survey periods using data from a recent range-wide meta-analysis. These estimates were a by-product of pair occupancy modeling but might provide insight into potential changes in the effect of the invasive barred owl on NSO detection rates. We used two-species multi-season occupancy models to estimate the probability of detection for NSOs on each of 11 study areas for each 2-week survey period and relative to the range-wide effect of barred owl presence or absence. Detection probabilities within the season generally increased from the earliest surveys in March through mid-season, decreasing again in the late season on five study areas. For three other study areas, detection rates were highest during the earliest survey periods in late March or early April. Estimates of cumulative seasonal detection of NSO (across a maximum of six within-season surveys) were less than 0.90 when barred owls (BO) were present on all but one study area, regardless of when surveys were conducted within a season. However, despite low detection rates, the probability that a territory was occupied when an NSO pair was not detected over six within-season surveys was also very low. When BO are not present on a territory, a six-survey protocol had a high probability of detecting an NSO pair at least once during the season on all study areas, except for the very lowest per-survey estimates. Conducting most surveys earlier in the season, when the probability of detecting pairs is highest (through May on most areas) could improve seasonal detection rates. However, alternative methods of population monitoring—such as the use of passive acoustic recorders—may be needed to continue monitoring NSO for research and management.

California, Oregon, Washington

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 sub-species 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 and 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.

Ecological Applications

Conspecific and congeneric interactions shape increasing rates of breeding dispersal of northern spotted owls

Breeding dispersal, the movement from one breeding territory to another, is rare for philopatric species that evolved within relatively stable environments, such as the old-growth coniferous forests of the Pacific Northwest. Although dispersal is not inherently maladaptive, the consequences of increased dispersal on population dynamics in populations whose historical dispersal rates are low could be significant, particularly for a declining species. We examined rates and possible causes of breeding dispersal based on a sample of 4,118 northern spotted owls ( Strix occidentalis caurina ) monitored in seven study areas over 28 yr, 1990–2017, in Oregon and Washington, USA. Using a multistate mark–resight analysis, we investigated the potential impacts of an emergent congeneric competitor (barred owl Strix varia ) and forest alteration (extrinsic factors), and social and individual conditions (intrinsic factors) on 408 successive and 1,372 nonsuccessive dispersal events between years. The annual probability of breeding dispersal increased for individual owls that had also dispersed in the previous year and decreased for owls on territories with historically high levels of reproduction. Intrinsic factors including pair status, prior reproductive success, and experience at a site, were also associated with breeding dispersal movements. The percent of monitored owls dispersing each year increased from ˜7% early in the study to ˜25% at the end of the study, which coincided with a rapid increase in numbers of invasive and competitively dominant barred owls. We suggest that the results presented here can inform spotted owl conservation efforts as we identify factors contributing to changing rates of demographic parameters including site fidelity and breeding dispersal. Our study further shows that increasing rates of breeding dispersal associated with population declines contribute to population instability and vulnerability of northern spotted owls to extinction, and the prognosis is unlikely to change unless active management interventions are undertaken.

California, Oregon, Washington

Range-wide declines of northern spotted owl populations in the Pacific Northwest: A meta-analysis

The northern spotted owl ( Strix occidentalis caurina ) inhabits older coniferous forests in the Pacific Northwest and has been at the center of forest management issues in this region. The immediate threats to this federally listed species include habitat loss and competition with barred owls ( Strix varia ), which invaded from eastern North America. We conducted a prospective meta-analysis to assess population trends and factors affecting those trends in northern spotted owls using 26 years of survey and capture-recapture data from 11 study areas across the owls' geographic range to analyze demographic traits, rates of population change, and occupancy parameters for spotted owl territories. We found that northern spotted owl populations experienced significant declines of 6–9% annually on 6 study areas and 2–5% annually on 5 other study areas. Annual declines translated to ≤35% of the populations remaining on 7 study areas since 1995. Barred owl presence on spotted owl territories was the primary factor negatively affecting apparent survival, recruitment, and ultimately, rates of population change. Analysis of spotted and barred owl detections in an occupancy framework corroborated the capture-recapture analyses with barred owl presence increasing territorial extinction and decreasing territorial colonization of spotted owls. While landscape habitat components reduced the effect of barred owls on these rates of decline, they did not reverse the negative trend. Our analyses indicated that northern spotted owl populations potentially face extirpation if the negative effects of barred owls are not ameliorated while maintaining northern spotted owl habitat across their range.

California, Oregon, Washington

Social status, forest disturbance, and Barred Owls shape long-term trends in breeding dispersal distance of Northern Spotted Owls

Dispersal among breeding sites in territorial animals (i.e. breeding dispersal) is driven by numerous selection pressures, including competition and spatiotemporal variation in habitat quality. The scale and trend of dispersal movements over time may signal changing conditions within the population or on the landscape. We examined 2,158 breeding dispersal events from 694 male and 608 female individually marked Northern Spotted Owls ( Strix occidentalis caurina ) monitored over 28 yr on 7 study areas to assess the relative importance of individual (sex, experience), reproductive (annual productivity, mate availability), and environmental (forest alteration, presence of competitor) sources of variation in breeding dispersal distance. Median breeding dispersal distance was 3.17 km, with 99% of all breeding dispersal events <37 km. Mean annual dispersal distances increased by 2.43 km in Oregon and 9.40 km in Washington between 1990 and 2017, which coincided with increases in annual detections of nonnative Barred Owl ( S. varia ). Frequency of breeding dispersal events, both among and within individuals, also increased over time. Female owls moved farther than males (median of 3.26 and 3.10 km, respectively), and birds with less experience (territory tenure) moved farther than those with more experience. Owls that were single in the year prior to dispersal moved 13–31% farther than those paired prior to dispersal. The greatest environmental change occurring over the course of our study was the expansion of Barred Owl populations. Breeding dispersal distance was positively related to Barred Owls in the study area and disturbance within the originating territory. While it appears that social factors continue to be important drivers of breeding dispersal distance in Spotted Owls, increased competition from Barred Owls and habitat alteration have a contributing effect. Increased breeding dispersal distances should be of concern for conservation efforts and considered in population monitoring because changing dispersal behavior may lead to higher rates of mortality and/or emigration from historical study areas.

Oregon, Washington

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).

Ecological Applications

The effects of habitat, climate, and Barred Owls on long-term demography of Northern Spotted Owls

Estimates of species' vital rates and an understanding of the factors affecting those parameters over time and space can provide crucial information for management and conservation. We used mark&ndash;recapture, reproductive output, and territory occupancy data collected during 1985&ndash;2013 to evaluate population processes of Northern Spotted Owls ( Strix occidentalis caurina ) in 11 study areas in Washington, Oregon, and northern California, USA. We estimated apparent survival, fecundity, recruitment, rate of population change, and local extinction and colonization rates, and investigated relationships between these parameters and the amount of suitable habitat, local and regional variation in meteorological conditions, and competition with Barred Owls ( Strix varia ). Data were analyzed for each area separately and in a meta-analysis of all areas combined, following a strict protocol for data collection, preparation, and analysis. We used mixed effects linear models for analyses of fecundity, Cormack-Jolly-Seber open population models for analyses of apparent annual survival (ϕ), and a reparameterization of the Jolly-Seber capture&ndash;recapture model (i.e. reverse Jolly-Seber; RJS) to estimate annual rates of population change (&lambda; RJS ) and recruitment. We also modeled territory occupancy dynamics of Northern Spotted Owls and Barred Owls in each study area using 2-species occupancy models. Estimated mean annual rates of population change (&lambda;) suggested that Spotted Owl populations declined from 1.2% to 8.4% per year depending on the study area. The weighted mean estimate of &lambda; for all study areas was 0.962 (&plusmn; 0.019 SE; 95% CI: 0.925&ndash;0.999), indicating an estimated range-wide decline of 3.8% per year from 1985 to 2013. Variation in recruitment rates across the range of the Spotted Owl was best explained by an interaction between total winter precipitation and mean minimum winter temperature. Thus, recruitment rates were highest when both total precipitation (29 cm) and minimum winter temperature (&minus;9.5&deg;C) were lowest. Barred Owl presence was associated with increased local extinction rates of Spotted Owl pairs for all 11 study areas. Habitat covariates were related to extinction rates for Spotted Owl pairs in 8 of 11 study areas, and a greater amount of suitable owl habitat was generally associated with decreased extinction rates. We observed negative effects of Barred Owl presence on colonization rates of Spotted Owl pairs in 5 of 11 study areas. The total amount of suitable Spotted Owl habitat was positively associated with colonization rates in 5 areas, and more habitat disturbance was associated with lower colonization rates in 2 areas. We observed strong declines in derived estimates of occupancy in all study areas. Mean fecundity of females was highest for adults (0.309 &plusmn; 0.027 SE), intermediate for 2-yr-olds (0.179 &plusmn; 0.040 SE), and lowest for 1-yr-olds (0.065 &plusmn; 0.022 SE). The presence of Barred Owls and habitat covariates explained little of the temporal variation in fecundity in most study areas. Climate covariates occurred in competitive fecundity models in 8 of 11 study areas, but support for these relationships was generally weak. The fecundity meta-analysis resulted in 6 competitive models, all of which included the additive effects of geographic region and annual time variation. The 2 top-ranked models also weakly supported the additive negative effects of the amount of suitable core area habitat, Barred Owl presence, and the amount of edge habitat on fecundity. We found strong support for a negative effect of Barred Owl presence on apparent survival of Spotted Owls in 10 of 11 study areas, but found few strong effects of habitat on survival at the study area scale. Climate covariates occurred in top or competitive survival models for 10 of 11 study areas, and in most cases the relationships were as predicted; however, there was little consistency among areas regarding the relative importance of specific climate covariates. In contrast, meta-analysis results suggested that Spotted Owl survival was higher across all study areas when the Pacific Decadal Oscillation (PDO) was in a warming phase and the Southern Oscillation Index (SOI) was negative, with a strongly negative SOI indicative of El Ni&ntilde;o events. The best model that included the Barred Owl covariate (BO) was ranked 4 th and also included the PDO covariate, but the BO effect was strongly negative. Our results indicated that Northern Spotted Owl populations were declining throughout the range of the subspecies and that annual rates of decline were accelerating in many areas. We observed strong evidence that Barred Owls negatively affected Spotted Owl populations, primarily by decreasing apparent survival and increasing local territory extinction rates. However, the amount of suitable owl habitat, local weather, and regional climatic patterns also were related to survival, occupancy (via colonization rate), recruitment, and, to a lesser extent, fecundity, although there was inconsistency in regard to which covariates were important for particular demographic parameters or across study areas. In the study areas where habitat was an important source of variation for Spotted Owl demographics, vital rates were generally positively associated with a greater amount of suitable owl habitat. However, Barred Owl densities may now be high enough across the range of the Northern Spotted Owl that, despite the continued management and conservation of suitable owl habitat on federal lands, the long-term prognosis for the persistence of Northern Spotted Owls may be in question without additional management intervention. Based on our study, the removal of Barred Owls from the Green Diamond Resources (GDR) study area had rapid, positive effects on Northern Spotted Owl survival and the rate of population change, supporting the hypothesis that, along with habitat conservation and management, Barred Owl removal may be able to slow or reverse Northern Spotted Owl population declines on at least a localized scale.

California, Oregon, Washington