Gail S. Olson

SMALL‐MAMMAL DENSITY ESTIMATION: A FIELD COMPARISON OF GRID‐BASED VS. WEB‐BASED DENSITY ESTIMATORS

Statistical models for estimating absolute densities of field populations of animals have been widely used over the last century in both scientific studies and wildlife management programs. To date, two general classes of density estimation models have been developed: models that use data sets from capture–recapture or removal sampling techniques (often derived from trapping grids) from which separate estimates of population size (N) and effective sampling area (Â) are used to calculate density (D = N/Â); and models applicable to sampling regimes using distance-sampling theory (typically transect lines or trapping webs) to estimate detection functions and densities directly from the distance data. However, few studies have evaluated these respective models for accuracy, precision, and bias on known field populations, and no studies have been conducted that compare the two approaches under controlled field conditions. In this study, we evaluated both classes of density estimators on known densities of e...

MODELING OF SITE OCCUPANCY DYNAMICS FOR NORTHERN SPOTTED OWLS, WITH EMPHASIS ON THE EFFECTS OF BARRED OWLS

Abstract Northern spotted owls (Strix occidentalis caurina) have been studied intensively since their listing as a threatened species by the U.S. Fish and Wildlife Service in 1990. Studies of spotted owl site occupancy have used various binary response measures, but most of these studies have made the assumption that detectability is perfect, or at least high and not variable. Further, previous studies did not consider temporal variation in site occupancy. We used relatively new methods for open population modeling of site occupancy that incorporated imperfect and variable detectability of spotted owls and allowed modeling of temporal variation in site occupancy, extinction, and colonization probabilities. We also examined the effects of barred owl (S. varia) presence on these parameters. We used spotted owl survey data from 1990 to 2002 for 3 study areas in Oregon, USA, and we used program MARK to develop and analyze site occupancy models. We found per visit detection probabilities averaged <0.70 and were highly variable among study years and study areas. Site occupancy probabilities for owl pairs declined greatly on 1 study area and slightly on the other 2 areas. For all owls, including singles and pairs, site occupancy was mostly stable through time. Barred owl presence had a negative effect on spotted owl detection probabilities, and it had either a positive effect on local-extinction probabilities or a negative effect on colonization probabilities. We conclude that further analyses of spotted owls must account for imperfect and variable detectability and barred owl presence to properly interpret results. Further, because barred owl presence is increasing within the range of northern spotted owls, we expect to see further declines in the proportion of sites occupied by spotted owls.

STATUS AND TRENDS IN DEMOGRAPHY OF NORTHERN SPOTTED OWLS, 1985-2003

Abstract We analyzed demographic data from northern spotted owls (Strix occidentalis caurina) from 14 study areas in Washington, Oregon, and California for 1985–2003. The purpose of our analyses was to provide an assessment of the status and trends of northern spotted owl populations throughout most of their geographic range. The 14 study areas made up approximately 12% of the range of the subspecies and included federal, tribal, private, and mixed federal and private lands. The study areas also included all the major forest types that the subspecies inhabits. The analyses followed rigorous protocols that were developed a priori and were the result of extensive discussions and consensus among the authors. Our primary objectives were to estimate fecundity, apparent survival (φ), and annual rate of population change (λ) and to determine if there were any temporal trends in these population parameters. In addition to analyses of data from individual study areas, we conducted 2 meta-analyses on each demographic parameter. One meta-analysis was conducted on all 14 areas, and the other was restricted to the 8 areas that constituted the Effectiveness Monitoring Plan for northern spotted owls under the Northwest Forest Plan. The average number of years of reproductive data per study area was 14 (range = 5–19), and the average number of recapture occasions per study area was 13 (range = 4–18). Only 1 study area had <12 years of data. Our results were based on 32,054 captures and resightings of 11,432 banded individuals for estimation of survival and 10,902 instances in which we documented the number of young produced by territorial females. The number of young fledged (NYF) per territorial female was analyzed by testing a suite of a priori models that included (1) effects of age, (2) linear or quadratic time trends, (3) presence of barred owls (Strix varia) in spotted owl territories, and (4) an even-odd year effect. The NYF varied among years on most study areas with a biennial cycle of high reproduction in even-numbered years and low reproduction in odd-numbered years. These cyclic fluctuations did not occur on all study areas, and the even–odd year effect waned during the last 5 years of the study. Fecundity was highest for adults (x̄ = 0.372, SE = 0.029), lower for 2-year-olds (x̄ = 0.208, SE = 0.032), and very low for 1-year-olds (x̄ = 0.074, SE = 0.029). Fecundity was stable over time for 6 areas (Rainier, Olympic, Warm Springs, H. J. Andrews, Klamath, and Marin), declining for 6 areas (Wenatchee, Cle Elum, Oregon Coast Range, Southern Oregon Cascades, Northwest California, and Simpson), and slightly increasing for 2 areas (Tyee, Hoopa). We found little association between NYF and the proportion of northern spotted owl territories where barred owls were detected, although results were suggestive of a negative effect of barred owls on the Wenatchee and Olympic study areas. The meta-analysis on fecundity indicated substantial annual variability with no increasing or decreasing trends. Fecundity was highest in the mixed-conifer region of eastern Washington (x̄ = 0.560, SE = 0.041) and lowest in the Douglas-fir (Pseudotsuga menziesii) region of the Oregon coast (x̄ = 0.306, SE = 0.039). We used Cormack–Jolly–Seber open population models and Program MARK to estimate apparent survival rates of owls >1 year old. We found no differences in apparent survival rates between sexes except for 1 area (Marin), which had only 6 years of data. Estimates of apparent survival from individual study areas indicated that there were differences among age classes with adults generally having higher survival than 1- and 2-year-olds. Apparent survival rates ranged from 0.750 (SE = 0.026) to 0.886 (SE = 0.010) for adults, 0.626 (SE = 0.073) to 0.886 (SE = 0.010) for 2-year-olds, and 0.415 (SE = 0.111) to 0.860 (SE = 0.017) for 1-year-olds. These estimates were comparable to survival rates from previous studies on the subspecies. We found evidence for negative time trends in survival rates on 5 study areas (Wenatchee, Cle Elum, Rainier, Olympic, and Northwest California) and no trends in survival on the remaining areas. There was evidence for negative effects of barred owls on apparent survival on 3 study areas (Wenatchee, Cle Elum, and Olympic). Survival rates of adult owls on the 8 Monitoring Areas generally were high, ranging from 0.85 (SE = 0.009) to 0.89 (SE = 0.010), but were declining on the Cle Elum, Olympic, and Northwestern California study areas. The meta-analysis of apparent survival indicated differences among regions and changes over time with a downward trend in the mixed-conifer and Douglas-fir regions of Washington. The meta-analysis of apparent survival also indicated that there was a negative association between fecundity and survival the following year, suggesting a cost of reproduction on survival. This effect was limited to the Douglas-fir and mixed-conifer regions of Washington and the Douglas-fir region of the Oregon Cascade Mountains. We used the reparameterized Jolly–Seber method (λRJS) to estimate annual rate of population change of territorial owls in the study areas. This estimate answers the question, Are these territorial owls being replaced in this geographically open population? Point estimates of λRJS were <1.0 for 12 of 13 study areas. The analyses provided strong evidence that populations on the Wenatchee, Cle Elum, Rainier, Olympic, Warm Springs, H. J. Andrews, Oregon Coast Ranges, and Simpson study areas were declining during the study. The mean λ̂RJS for the 13 study areas was 0.963 (SE = 0.009), suggesting that populations over all the areas were declining about 3.7% per year during the study. The mean λ̂RJS for the 8 monitoring areas for the Northwest Forest Plan was 0.976 (SE = 0.007) compared to a mean of 0.942 (SE = 0.016) for the other study areas, a 2.4-versus-5.8% decline per year. This suggested that owl populations on federal lands had higher demographic rates than elsewhere; thus, the Northwest Forest Plan appeared to have a positive effect on demography of northern spotted owls. Populations were doing poorest in Washington, where apparent survival rates and populations were declining on all 4 study areas. Our estimates of λRJS were generally lower than those reported in a previous analysis (λ̂RJS = 0.997, SE = 0.003) for many of the same areas at an earlier date. The possible causes of population declines include but are not limited to habitat loss from timber harvest and fires, competition with barred owls, and weather patterns.

MODELING DEMOGRAPHIC PERFORMANCE OF NORTHERN SPOTTED OWLS RELATIVE TO FOREST HABITAT IN OREGON

Abstract Northern spotted owls (Strix occidentalis caurina) are known to be associated with late-successional forests in the Pacific Northwest of the United States, but the effects of habitat on their demographic performance are relatively unknown. We developed statistical models relating owl survival and productivity to forest cover types within the Roseburg Study Area in the Oregon Coast Range of Oregon, USA. We further combined these demographic parameters using a Leslie-type matrix to obtain an estimate of habitat fitness potential for each owl territory (n = 94). We used mark–recapture methods to develop models for survival and linear mixed models for productivity. We measured forest composition and landscape patterns at 3 landscape scales centered on nest and activity sites within owl territories using an aerial photo-based map and a Geographic Information System (GIS). We also considered additional covariates such as age, sex, and presence of barred owls (Strix varia), and seasonal climate variables (temperature and precipitation) in our models. We used Akaikes Information Criterion (AIC) to rank and compare models. Survival had a quadratic relationship with the amount of late- and mid-seral forests within 1,500 m of nesting centers. Survival also was influenced by the amount of precipitation during the nesting season. Only 16% of the variability in survival was accounted for by our best model, but 85% of this was due to the habitat variable. Reproductive rates fluctuated biennially and were positively related to the amount of edge between late- and mid-seral forests and other habitat classes. Reproductive rates also were influenced by parent age, amount of precipitation during nesting season, and presence of barred owls. Our best model accounted for 84% of the variability in productivity, but only 3% of that was due to the habitat variable. Estimates of habitat fitness potential (which may range from 0 to infinity) for the 94 territories ranged from 0.74 to 1.15 (x̄ = 1.05, SE = 0.07). All but 1 territory had 95% confidence intervals overlapping 1.0, indicating a potentially stable population based on habitat pattern. Our results seem to indicate that while mid- and late-seral forests are important to owls, a mixture of these forest types with younger forest and nonforest may be best for owl survival and reproduction. Our results are consistent with those of researchers in northern California, USA, who used similar methods in their analyses. However, we believe that given the low variability in survival and productivity attributed to habitat, further study is needed to confirm our conclusions before they can be used to guide forest management actions for spotted owls.

THE RELATIONSHIP BETWEEN HABITAT CHARACTERISTICS AND DEMOGRAPHIC PERFORMANCE OF NORTHERN SPOTTED OWLS IN SOUTHERN OREGON

Abstract We used data from Northern Spotted Owl (Strix occidentalis caurina) territories to model the effects of habitat (particularly intermediate-aged forest stand types), climate, and nonhabitat covariates (i.e., age, sex) on owl reproductive rate and apparent survival in southwestern Oregon. Our best model for reproductive rate included an interaction between a cyclic, annual time trend and male breeding experience, with higher reproductive rates in even years compared to odd, particularly for males with previous breeding experience. Reproductive rate was also negatively related to the amount of winter precipitation and positively related to the proportion of old-growth forest near the owl territory center. Apparent survival was not associated with age, sex, climate or any of the intermediate-aged forest types, but was positively associated with the proportion of older forest near the territory center in a pseudothreshold pattern. The quadratic structure of the proportion of nonhabitat farther from the nest or primary roost site was also part of our best survival model. Survival decreased dramatically when the amount of nonhabitat exceeded ∼50%. Habitat fitness potential estimates (λ̂h) for 97 owl territories ranged from 0.29–1.09, with a mean of 0.86 ± 0.02. Owl territories with habitat fitness potentials <1.0 were generally characterized by <40%–50% old forest habitat near the territory center. Our results indicate that both apparent survival and reproductive rate are positively associated with older forest types close to the nest or primary roost site. We found no support for either a positive or negative direct effect of intermediate-aged forests on either survival or reproductive rate.

Effects of drought and prolonged winter on Townsend's ground squirrel demography in shrubsteppe habitats

During a mark–recapture study of Townsend’s ground squirrels (Spermophilus townsendii) on 20 sites in the Snake River Birds of Prey National Conservation Area, Idaho, in 1991 through 1994, 4407 animals were marked in 17639 capture events. This study of differences in population dynamics of Townsend’s ground squirrels among habitats spanned a drought near the extreme of the 130-yr record, followed by prolonged winter conditions. Townsend’s ground squirrels have a short active season (≈4 mo) in which to reproduce and store fat for overwintering. Their food consists largely of succulent grasses and forbs in this dry shrubsteppe and grassland habitat. The drought in the latter half of the 1992 active season produced early drying of Sandberg’s bluegrass (Poa secunda) and was associated with low adult and juvenile body masses prior to immergence into estivation/hibernation. The following prolonged winter was associated with late emergence of females in 1993. Early-season body masses of adults were low in 1993 r...

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

ABSTRACT 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.

Local weather, regional climate, and annual survival of the northern spotted owl

Abstract. We used an information-theoretical approach and Cormack—Jolly—Seber models for open populations in program MARK to examine relationships between survival rates of Northern Spotted Owls and a variety of local weather variables and long-term climate variables. In four of the six populations examined, survival was positively associated with wetter than normal conditions during the growing season or high summer temperatures. At the three study areas located at the highest elevations, survival was positively associated with winter temperature but also had a negative or quadratic relation with the number of storms and winter precipitation. A meta-analysis of all six areas combined indicated that annual survival was most strongly associated with phase shifts in the Southern Oscillation and Pacific Decadal Oscillation, which reflect large-scale temperature and precipitation patterns in this region. Climate accounted for a variable amount (1–41%) of the total process variation in annual survival but for more year-to-year variation (3–66%) than did spatial variation among owl territories (0–7%). Negative associations between survival and cold, wet winters and nesting seasons were similar to those found in other studies of the Spotted Owl. The relationships between survival and growing-season precipitation and regional climate patterns, however, had not been reported for this species previously. Climate-change models for the first half of the 21st century predict warmer, wetter winters and hotter, drier summers for the Pacific Northwest. Our results indicate that these conditions could decrease Spotted Owl survival in some areas.

Barred Owls and Landscape Attributes Influence Territory Occupancy of Northern Spotted Owls

We used multi-season occupancy analyses to model 2 fates of northern spotted owl territories in relation to habitat amount, habitat fragmentation, and the presence of barred owls in Washington State, USA, 1989–2005. Local colonization is the probability a territory unoccupied by a spotted owl in year i would be occupied in year i + 1, and local extinction is the probability a territory that was occupied by a spotted owl in year i would be unoccupied in year i + 1. We found a negative relationship between local extinction probability and amount of late-seral forest edge. We found a negative relationship between colonization probability and the number of late-seral forest patches (higher fragmentation), and a negative relationship between colonization probability and the amount of non-habitat within 600 m of a spotted owl territory center (Akaike weight = 0.59). The presence of barred owls was positively related to extinction probability and negatively related to detection probability of spotted owls. The negative relationship between presence of barred owls and detectability of spotted owls indicated that spotted owls could be modifying their calling behavior in the presence of barred owls. The positive relationship between barred owl detections and local extinction probability suggests that because of competition with barred owls, spotted owls are being displaced. Published 2014. This article is a U.S. Government work and is in the public domain in the USA.