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Gary C. White

Publications and source records attributed to Gary C. White.

At least 19 recordsLinked to original sources

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↗

Invader removal triggers competitive release in a threatened avian predator

Invasive species can cause extinctions of native species and widespread biodiversity loss. Invader removal is a common management response, but the use of long-term field experiments to characterize effectiveness of removals in benefitting impacted native species is rare. We used a large-scale removal experiment to investigate the demographic response of a threatened native species, the northern spotted owl, to removal of an invasive competitor species, the barred owl. Removal of barred owls had a strong, positive effect on survival of spotted owls, which arrested long-term population declines of spotted owls. The results demonstrate that the long-term persistence of spotted owls will depend heavily on reducing the negative impacts of barred owls while simultaneously addressing other threats, such as habitat loss.

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↗

A multistate open robust design: population dynamics, reproductive effort, and phenology of sea turtles from tagging data

Understanding population dynamics, and how it is influenced by exogenous and endogenous factors, is important to the study and conservation of species. Moreover, for migratory species, the phenology and duration of use of a given location can also influence population structure and dynamics. For many species, breeding abundance, survival, and reproductive performance, as well as phenology of nesting, are often the most accessible, and therefore practical, elements of their life history to study. For a population of hawksbill sea turtles (Eretmochelys imbricata), we modeled population change for nesters and total adult females, survival, and breeding probability, from 25 years of intensive tagging data. We modeled breeding probability as a function of the number of years since last breeding, and tested for differences between neophyte and experienced nesters. For each year, we also estimated the number of clutches deposited per female, and phenology of use, for neophytes and experienced nesters. In order to implement the analysis we developed a novel generalized multistate open robust design mark-recapture modeling framework, with parameters for survival and transition probabilities, and for each primary period, state structure and arrival, persistence, and detection probabilities. Derived parameters included abundance of observable and unobservable components of the population, residence time, expected arrival and departure periods, and per-period intensity of study area use. Abundance of nesters increased over most of the time series. Survival probability was 0.935 (se = 0.01). All hawksbills skipped at least one year of nesting. Breeding probability increased by skipping a second year, but then decreased thereafter. Subsequent breeding probability was lower for neophyte nesters than for experienced nesters, but the effect was weaker than the effect of years since breeding. Clutch frequency varied by year, with no discernable pattern of differences between neophytes and experienced nesters. Mean arrival and departure dates also varied, with a slight shift of nesting activity to earlier in the season. The multistate open robust design model developed here provides a flexible framework for modeling the dynamics of structured migratory populations, and the phenology and duration of their seasonal use of study areas.

Caribbean, Jumby Bay Leeward Islands, Long Island↗

Overwintering strategies of migratory birds: a novel approach for estimating seasonal movement patterns of residents and transients

Our understanding of movement patterns in wildlife populations has played an important role in current ecological knowledge and can inform landscape conservation decisions. Direct measures of movement can be obtained using marked individuals, but this requires tracking individuals across a landscape or multiple sites. We demonstrate how movements can be estimated indirectly using single-site, capture–mark–recapture (CMR) data with a multi-state open robust design with state uncertainty model (MSORD-SU). We treat residence and transience as two phenotypic states of overwintering migrants and use time- and state-dependent probabilities of site entry and persistence as indirect measures of movement. We applied the MSORD-SU to data on eight species of overwintering Neotropical birds collected in 14 countries between 2002 and 2011. In addition to entry and persistence probabilities, we estimated the proportions of residents at a study site and mean residence times. We identified overwintering movement patterns and residence times that contrasted with prior categorizations of territoriality. Most species showed an evidence of residents entering sites at multiple time intervals, with transients tending to enter between peak resident movement times. Persistence and the proportion of residents varied by latitude, but were not always positively correlated for a given species. Synthesis and applications . Our results suggest that migratory songbirds commonly move among habitats during the overwintering period. Substantial proportions of populations appear to be comprised of transient individuals, and residents tend to persist at specific sites for relatively short periods of time. This information on persistence and movement patterns should be explored for specific habitats to guide landscape management on the wintering grounds, such as determining which habitats are conserved or restored as part of certification programmes of tropical agroforestry crops. We suggest that research and conservation efforts on Neotropical migrant songbirds focus on identifying landscape configurations and regional habitat networks that support these diverse overwintering strategies to secure full life cycle conservation.

Journal of Applied Ecology↗

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–recapture, reproductive output, and territory occupancy data collected during 1985–2013 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–recapture model (i.e. reverse Jolly-Seber; RJS) to estimate annual rates of population change (λ 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 (λ) suggested that Spotted Owl populations declined from 1.2% to 8.4% per year depending on the study area. The weighted mean estimate of λ for all study areas was 0.962 (± 0.019 SE; 95% CI: 0.925–0.999), 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 (−9.5°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 ± 0.027 SE), intermediate for 2-yr-olds (0.179 ± 0.040 SE), and lowest for 1-yr-olds (0.065 ± 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 Niño 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↗

Density dependence, whitebark pine, and vital rates of grizzly bears

Understanding factors influencing changes in population trajectory is important for effective wildlife management, particularly for populations of conservation concern. Annual population growth of the grizzly bear ( Ursus arctos ) population in the Greater Yellowstone Ecosystem, USA has slowed from 4.2–7.6% during 1983–2001 to 0.3–2.2% during 2002–2011. Substantial changes in availability of a key food source and bear population density have occurred. Whitebark pine ( Pinus albicaulis ), the seeds of which are a valuable but variable fall food for grizzly bears, has experienced substantial mortality primarily due to a mountain pine beetle ( Dendroctonus ponderosae ) outbreak that started in the early 2000s. Positive growth rates of grizzly bears have resulted in populations reaching high densities in some areas and have contributed to continued range expansion. We tested research hypotheses to examine if changes in vital rates detected during the past decade were more associated with whitebark pine decline or, alternatively, increasing grizzly bear density. We focused our assessment on known-fate data to estimate survival of cubs-of-the-year (cubs), yearlings, and independent bears (≥2 yrs), and reproductive transition of females from having no offspring to having cubs. We used spatially and temporally explicit indices for grizzly bear density and whitebark pine mortality as individual covariates. Models indicated moderate support for an increase in survival of independent male bears over 1983–2012, whereas independent female survival did not change. Cub survival, yearling survival, and reproductive transition from no offspring to cubs all changed during the 30-year study period, with lower rates evident during the last 10–15 years. Cub survival and reproductive transition were negatively associated with an index of grizzly bear density, indicating greater declines where bear densities were higher. Our analyses did not support a similar relationship for the index of whitebark pine mortality. The results of our study support the interpretation that slowing of population growth during the last decade was associated more with increasing grizzly bear density than the decline in whitebark pine. Grizzly bear density and its potential effect on vital rates and population trajectory warrant consideration for management of the grizzly bear population in the Greater Yellowstone Ecosystem.

Idaho, Montana, Wyoming↗

Re-evaluation of Yellowstone grizzly bear population dynamics not supported by empirical data: response to Doak & Cutler

Doak and Cutler critiqued methods used by the Interagency Grizzly Bear Study Team (IGBST) to estimate grizzly bear population size and trend in the Greater Yellowstone Ecosystem. Here, we focus on the premise, implementation, and interpretation of simulations they used to support their arguments. They argued that population increases documented by IGBST based on females with cubs-of-the-year were an artifact of increased search effort. However, we demonstrate their simulations were neither reflective of the true observation process nor did their results provide statistical support for their conclusion. They further argued that survival and reproductive senescence should be incorporated into population projections, but we demonstrate their choice of extreme mortality risk beyond age 20 and incompatible baseline fecundity led to erroneous conclusions. The conclusions of Doak and Cutler are unsubstantiated when placed within the context of a thorough understanding of the data, study system, and previous research findings and publications.

Conservation Letters↗

Increased flexibility for modeling telemetry and nest-survival data using the multistate framework

Although telemetry is one of the most common tools used in the study of wildlife, advances in the analysis of telemetry data have lagged compared to progress in the development of telemetry devices. We demonstrate how standard known-fate telemetry and related nest-survival data analysis models are special cases of the more general multistate framework. We present a short theoretical development, and 2 case examples regarding the American black duck and the mallard. We also present a more complex lynx data analysis. Although not necessary in all situations, the multistate framework provides additional flexibility to analyze telemetry data, which may help analysts and biologists better deal with the vagaries of real-world data collection.

Journal of Wildlife Management↗

Insights into the latent multinomial model through mark-resight data on female grizzly bears with cubs-of-the-year

Mark-resight designs for estimation of population abundance are common and attractive to researchers. However, inference from such designs is very limited when faced with sparse data, either from a low number of marked animals, a low probability of detection, or both. In the Greater Yellowstone Ecosystem, yearly mark-resight data are collected for female grizzly bears with cubs-of-the-year (FCOY), and inference suffers from both limitations. To overcome difficulties due to sparseness, we assume homogeneity in sighting probabilities over 16 years of bi-annual aerial surveys. We model counts of marked and unmarked animals as multinomial random variables, using the capture frequencies of marked animals for inference about the latent multinomial frequencies for unmarked animals. We discuss undesirable behavior of the commonly used discrete uniform prior distribution on the population size parameter and provide OpenBUGS code for fitting such models. The application provides valuable insights into subtleties of implementing Bayesian inference for latent multinomial models. We tie the discussion to our application, though the insights are broadly useful for applications of the latent multinomial model.

Journal of Agricultural, Biological, and Environme↗

Estimating parameters of hidden Markov models based on marked individuals: use of robust design data

Development and use of multistate mark-recapture models, which provide estimates of parameters of Markov processes in the face of imperfect detection, have become common over the last twenty years. Recently, estimating parameters of hidden Markov models, where the state of an individual can be uncertain even when it is detected, has received attention. Previous work has shown that ignoring state uncertainty biases estimates of survival and state transition probabilities, thereby reducing the power to detect effects. Efforts to adjust for state uncertainty have included special cases and a general framework for a single sample per period of interest. We provide a flexible framework for adjusting for state uncertainty in multistate models, while utilizing multiple sampling occasions per period of interest to increase precision and remove parameter redundancy. These models also produce direct estimates of state structure for each primary period, even for the case where there is just one sampling occasion. We apply our model to expected value data, and to data from a study of Florida manatees, to provide examples of the improvement in precision due to secondary capture occasions. We also provide user-friendly software to implement these models. This general framework could also be used by practitioners to consider constrained models of particular interest, or model the relationship between within-primary period parameters (e.g., state structure) and between-primary period parameters (e.g., state transition probabilities).

Ecology↗

Targeted surveillance for highly pathogenic avian influenza in migratory waterfowl across the conterminous United States: chapter 12

Introduction of Asian strain H5N1 Highly Pathogenic avian influenca via waterfowl migration is one potential route of entry into the United States. In conjunction with state, tribe, and laboratory partners, the United States Department of Agriculture collected and tested 124,603 wild bird samples in 2006 as part of a national surveillance effort. A sampling plan was devised to increase the probability fo detecting Asian strain H5N1 at a national scale. Band recovery data were used to identify and prioritize sampling for wild migratory waterfowl, resulting in spatially targeted sampling recommendations focused on reads with high numbers of recoveries. We also compared the spatial and temporal distribution of the 2006 cloacal and fecal waterfowl sampling effort to the bird banding recovery data and found concordance between the two .Finally, we present improvements made to the 2007 fecal sampling component of the surveillance plan and suggest further improvements for future sampling.

Book chapter↗

Response of brown treesnakes to reduction of their rodent prey

Trapping brown treesnakes (Boiga irregularis; BTS) with live-mouse (Mus domesticus) lures is the principal control technique for this invasive species on Guam. Lure-based trapping is also used on other islands as a precaution against undetected arrivals and in response to verified BTS sightings. However, the effectiveness of lure-based trapping on other islands is questionable, as it has yielded no BTS despite other evidence of their presence. Some evidence suggests that high rodent numbers may interfere with BTS control. To test the relationship between rodent abundance and snake trappability, we conducted a controlled, replicated field experiment incorporating a rodenticide treatment during a BTS mark-recapture study. Using open population modeling in Program MARK, we estimated BTS apparent survival and recapture probabilities. Rodent reduction increased BTS recapture probabilities by 52-65% in 2002 and 22-36% in 2003, and it decreased apparent survival by <1% both years. This appears to be the first published instance of manipulating wild prey to influence snake behavior. Rodent reduction may enhance detection and control of BTS with traps on Guam and other islands. It may also amplify the effectiveness of oral toxicants against BTS.

Journal of Wildlife Management↗

Temporal patterns of apparent leg band retention in North American geese

An important assumption of mark?recapture studies is that individuals retain their marks, which has not been assessed for goose reward bands. We estimated aluminum leg band retention probabilities and modeled how band retention varied with band type (standard vs. reward band), band age (1-40 months), and goose characteristics (species and size class) for Canada (Branta canadensis), cackling (Branta hutchinsii), snow (Chen caerulescens), and Ross?s (Chen rossii) geese that field coordinators double-leg banded during a North American goose reward band study (N = 40,999 individuals from 15 populations). We conditioned all models in this analysis on geese that were encountered with >1 leg band still attached (n = 5,747 dead recoveries and live recaptures). Retention probabilities for standard aluminum leg bands were high (estimate of 0.9995, SE = 0.001) and constant over 1-40 months. In contrast, apparent retention probabilities for reward bands demonstrated an interactive relationship between 5 size and species classes (small cackling, medium Canada, large Canada, snow, and Ross?s geese). In addition, apparent retention probabilities for each of the 5 classes varied quadratically with time, being lower immediately after banding and at older age classes. The differential retention probabilities among band type (reward vs. standard) that we observed suggests that 1) models estimating reporting probability should incorporate differential band loss if it is nontrivial, 2) goose managers should consider the costs and benefits of double-banding geese on an operational basis, and 3) the United States Geological Survey Bird Banding Lab should modify protocols for receiving recovery data.

Journal of Wildlife Management↗

Factors influencing reporting and harvest probabilities in North American geese

We assessed variation in reporting probabilities of standard bands among species, populations, harvest locations, and size classes of North American geese to enable estimation of unbiased harvest probabilities. We included reward (US10,20,30,50, or100) and control (0) banded geese from 16 recognized goose populations of 4 species: Canada (Branta canadensis), cackling (B. hutchinsii), Ross's (Chen rossii), and snow geese (C. caerulescens). We incorporated spatially explicit direct recoveries and live recaptures into a multinomial model to estimate reporting, harvest, and band-retention probabilities. We compared various models for estimating harvest probabilities at country (United States vs. Canada), flyway (5 administrative regions), and harvest area (i.e., flyways divided into northern and southern sections) scales. Mean reporting probability of standard bands was 0.73 (95 CI 0.690.77). Point estimates of reporting probabilities for goose populations or spatial units varied from 0.52 to 0.93, but confidence intervals for individual estimates overlapped and model selection indicated that models with species, population, or spatial effects were less parsimonious than those without these effects. Our estimates were similar to recently reported estimates for mallards (Anas platyrhynchos). We provide current harvest probability estimates for these populations using our direct measures of reporting probability, improving the accuracy of previous estimates obtained from recovery probabilities alone. Goose managers and researchers throughout North America can use our reporting probabilities to correct recovery probabilities estimated from standard banding operations for deriving spatially explicit harvest probabilities.

Journal of Wildlife Management↗

Demography and genetic structure of a recovering grizzly bear population

Grizzly bears (brown bears; Ursus arctos) are imperiled in the southern extent of their range worldwide. The threatened population in northwestern Montana, USA, has been managed for recovery since 1975; yet, no rigorous data were available to monitor program success. We used data from a large noninvasive genetic sampling effort conducted in 2004 and 33 years of physical captures to assess abundance, distribution, and genetic health of this population. We combined data from our 3 sampling methods (hair trap, bear rub, and physical capture) to construct individual bear encounter histories for use in Huggins-Pledger closed mark-recapture models. Our population estimate, N?? = 765 (95% CI = 715-831) was more than double the existing estimate derived from sightings of females with young. Based on our results, the estimated known, human-caused mortality rate in 2004 was 4.6% (95% CI = 4.2-4.9%), slightly above the 4% considered sustainable; however, the high proportion of female mortalities raises concern. We used location data from telemetry, confirmed sightings, and genetic sampling to estimate occupied habitat. We found that grizzly bears occupied 33,480 km2 in the Northern Continental Divide Ecosystem (NCDE) during 1994-2007, including 10,340 km beyond the Recovery Zone. We used factorial correspondence analysis to identify potential barriers to gene flow within this population. Our results suggested that genetic interchange recently increased in areas with low gene flow in the past; however, we also detected evidence of incipient fragmentation across the major transportation corridor in this ecosystem. Our results suggest that the NCDE population is faring better than previously thought, and they highlight the need for a more rigorous monitoring program.

Journal of Wildlife Management↗

Population dynamics of spotted owls in the Sierra Nevada, California

The California spotted owl (Strix occidentalis occidentalis) is the only spotted owl subspecies not listed as threatened or endangered under the United States Endangered Species Act despite petitions to list it as threatened. We conducted a meta-analysis of population data for 4 populations in the southern Cascades and Sierra Nevada, California, USA, from 1990 to 2005 to assist a listing evaluation by the United States Fish and Wildlife Service. Our study areas (from N to S) were on the Lassen National Forest (LAS), Eldorado National Forest (ELD), Sierra National Forest (SIE), and Sequoia and Kings Canyon National Parks (SKC). These study areas represented a broad spectrum of habitat and management conditions in these mountain ranges. We estimated apparent survival probability, reproductive output, and rate of population change for spotted owls on individual study areas and for all study areas combined (meta-analysis) using model selection or model-averaging based on maximum-likelihood estimation. We followed a formal protocol to conduct this analysis that was similar to other spotted owl meta-analyses. Consistency of field and analytical methods among our studies reduced confounding methodological effects when evaluating results. We used 991 marked spotted owls in the analysis of apparent survival. Apparent survival probability was higher for adult than for subadult owls. There was little difference in apparent survival between male and female owls. Model-averaged mean estimates of apparent survival probability of adult owls varied from 0.811 ?? 0.021 for females at LAS to 0.890 ?? 0.016 for males at SKC. Apparent survival increased over time for owls of all age classes at LAS and SIE, for adults at ELD, and for second-year subadults and adults at SKC. The meta-analysis of apparent survival, which included only adult owls, confirmed an increasing trend in survival over time. Survival rates were higher for owls on SKC than on the other study areas. We analyzed data from 1,865 observations of reproductive outcomes for female spotted owls. The proportion of subadult females among all territorial females of known age ranged from 0.00 to 0.25 among study areas and years. The proportion of subadults among female spotted owls was negatively related to reproductive output (no. of young fledged/territorial F owl) for ELD and SIE. Eldorado study area and LAS showed an alternate-year trend in reproductive output, with higher output in even-numbered years. Mean annual reproductive output was 0.988 ?? 0.154 for ELD, 0.624 ?? 0.140 for LAS, 0.478 ?? 0.106 for SIE, and 0.555 ?? 0.110 for SKC. Eldorado Study Area exhibited a declining trend and the greatest variation in reproductive output over time, whereas SIE and SKC, which had the lowest reproductive output, had the lowest temporal variation. Meta-analysis confirmed that reproductive output varied among study areas. Reproductive output was highest for adults, followed by second-year subadults, and then by first-year subadults. We used 842 marked subadult and adult owls to estimate population rate of change. Modeling indicated that ??t (??t is the finite rate of population change estimated using the reparameterized JollySeber estimator Pradel 1996) was either stationary (LAS and SIE) or increasing after an initial decrease (ELD and SKC). Mean estimated ??t for the 4 study areas was 1.007 (95 CI 0.9521.066) for ELD; 0.973 (95 CI 0.9461.001) for LAS; 0.992 (95 CI 0.9661.018) for SIE; and 1.006 (95 CI 0.9471.068) for SKC. The best meta-analysis model of population trend indicated that ?? varied across time but was similar in trend among the study areas. Our estimates of realized population change (??t; Franklin et al. 2004), which we estimated as the product 1 ?? ??3 ?? ??4 ?? .?? ??k -1, were based on estimates of ??t from individual study areas and did not require estimating annual population size for each study area. Trends represented the proportion of the population size in the first ye

Wildlife Monographs↗

Hazards affecting grizzly bear survival in the Greater Yellowstone Ecosystem

During the past 2 decades, the grizzly bear ( Ursus arctos ) population in the Greater Yellowstone Ecosystem (GYE) has increased in numbers and expanded its range. Early efforts to model grizzly bear mortality were principally focused within the United States Fish and Wildlife Service Grizzly Bear Recovery Zone, which currently represents only about 61% of known bear distribution in the GYE. A more recent analysis that explored one spatial covariate that encompassed the entire GYE suggested that grizzly bear survival was highest in Yellowstone National Park, followed by areas in the grizzly bear Recovery Zone outside the park, and lowest outside the Recovery Zone. Although management differences within these areas partially explained differences in grizzly bear survival, these simple spatial covariates did not capture site-specific reasons why bears die at higher rates outside the Recovery Zone. Here, we model annual survival of grizzly bears in the GYE to 1) identify landscape features (i.e., foods, land management policies, or human disturbances factors) that best describe spatial heterogeneity among bear mortalities, 2) spatially depict the differences in grizzly bear survival across the GYE, and 3) demonstrate how our spatially explicit model of survival can be linked with demographic parameters to identify source and sink habitats. We used recent data from radiomarked bears to estimate survival (1983–2003) using the known-fate data type in Program MARK. Our top models suggested that survival of independent (age ≥2 yr) grizzly bears was best explained by the level of human development of the landscape within the home ranges of bears. Survival improved as secure habitat and elevation increased but declined as road density, number of homes, and site developments increased. Bears living in areas open to fall ungulate hunting suffered higher rates of mortality than bears living in areas closed to hunting. Our top model strongly supported previous research that identified roads and developed sites as hazards to grizzly bear survival. We also demonstrated that rural homes and ungulate hunting negatively affected survival, both new findings. We illustrate how our survival model, when linked with estimates of reproduction and survival of dependent young, can be used to identify demographically the source and sink habitats in the GYE. Finally, we discuss how this demographic model constitutes one component of a habitat-based framework for grizzly bear conservation. Such a framework can spatially depict the areas of risk in otherwise good habitat, providing a focus for resource management in the GYE.

Idaho, Montana, Wyoming↗