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M.A. Haroldson

Publications and source records attributed to M.A. Haroldson.

11 recordsLinked to original sources

Population fragmentation and inter-ecosystem movements of grizzly bears in Western Canada and the Northern United States

Population fragmentation compromises population viability, reduces a species ability to respond to climate change, and ultimately may reduce biodiversity. We studied the current state and potential causes of fragmentation in grizzly bears over approximately 1,000,000 km 2 of western Canada, the northern United States (US), and southeast Alaska. We compiled much of our data from projects undertaken with a variety of research objectives including population estimation and trend, landscape fragmentation, habitat selection, vital rates, and response to human development. Our primary analytical techniques stemmed from genetic analysis of 3,134 bears, supplemented with radiotelemetry data from 792 bears. We used 15 locus microsatellite data coupled withmeasures of genetic distance, isolation-by-distance (IBD) analysis, analysis of covariance (ANCOVA), linear multiple regression, multi-factorial correspondence analysis (to identify population divisions or fractures with no a priori assumption of group membership), and population-assignment methods to detect individual migrants between immediately adjacent areas. These data corroborated observations of inter-area movements from our telemetry database. In northern areas, we found a spatial genetic pattern of IBD, although there was evidence of natural fragmentation from the rugged heavily glaciated coast mountains of British Columbia (BC) and the Yukon. These results contrasted with the spatial pattern of fragmentation in more southern parts of their distribution. Near the Canada-US border area, we found extensive fragmentation that corresponded to settled mountain valleys andmajor highways. Genetic distances across developed valleys were elevated relative to those across undeveloped valleys in central and northern BC. In disturbed areas, most inter-area movements detected were made by male bears, with few female migrants identified. North-south movements within mountain ranges (Mts) and across BC Highway 3 were more common than east-west movements across settled mountain valleys separating Mts. Our results suggest that relatively distinct subpopulations exist in this region, including the Cabinet, Selkirk South, and the decadesisolated Yellowstone populations. Current movement rates do not appear sufficient to consider the subpopulations we identify along the Canada-US border as 1 inter-breeding unit. Although we detected enough male movement to mediate gene flow, the current low rate of female movement detected among areas is insufficient to provide a demographic rescue effect between areas in the immediate future (0-15 yr). In Alberta, we found fragmentation corresponded to major east-west highways (Highways 3, 11, 16, and 43) and most inter-area movements were made by males. Gene flow and movement rates between Alberta and BC were highest across the Continental Divide south of Highway 1 and north of Highway 16. In the central region between Highways 1 and 11, we found evidence of natural fragmentation associated with the extensive glaciers and icefields along the Continental Divide. The discontinuities that we identified would form appropriate boundaries formanagement units. We related sex-specific movement rates between adjacent areas to several metrics of human use (highway traffic, settlement, and humancaused mortality) to understand the causes of fragmentation. This analysis used data from 1,508 bears sampled over a 161,500-km 2 area in southeastern BC, western Alberta, northern Idaho, and northern Montana during 1979-2007. This area was bisected by numerous human transportation and settlement corridors of varying intensity and complexity. We used multiple linear regression and ANCOVA to document the responses of female and male bears to disturbance. Males and females both demonstrated reduced movement rates with increasing settlement and traffic. However, females reduced their movement rates dramatically when settlement increased to >20% of the fracture zone. At this same

Wildlife Monographs

Grizzly bear population vital rates and trend in the Northern Continental Divide Ecosystem, Montana

We estimated grizzly bear ( Ursus arctos ) population vital rates and trend for the Northern Continental Divide Ecosystem (NCDE), Montana, between 2004 and 2009 by following radio-collared females and observing their fate and reproductive performance. Our estimates of dependent cub and yearling survival were 0.612 (95% CI = 0.300–0.818) and 0.682 (95% CI = 0.258–0.898). Our estimates of subadult and adult female survival were 0.852 (95% CI = 0.628–0.951) and 0.952 (95% CI = 0.892–0.980). From visual observations, we estimated a mean litter size of 2.00 cubs/litter. Accounting for cub mortality prior to the first observations of litters in spring, our adjusted mean litter size was 2.27 cubs/litter. We estimated the probabilities of females transitioning from one reproductive state to another between years. Using the stable state probability of 0.322 (95% CI = 0.262–0.382) for females with cub litters, our adjusted fecundity estimate ( m x ) was 0.367 (95% CI = 0.273–0.461). Using our derived rates, we estimated that the population grew at a mean annual rate of approximately 3% (λ = 1.0306, 95% CI = 0.928–1.102), and 71.5% of 10,000 Monte Carlo simulations produced estimates of λ > 1.0. Our results indicate an increasing population trend of grizzly bears in the NCDE. Coupled with concurrent studies of population size, we estimate that over 1,000 grizzly bears reside in and adjacent to this recovery area. We suggest that monitoring of population trend and other vital rates using radioed females be continued.

Montana

Cougar survival and source-sink structure on Greater Yellowstone's Northern Range

We studied survival and causes of mortality of radiocollared cougars ( Puma concolor ) on the Greater Yellowstone Northern Range (GYNR) prior to (1987–1994) and after wolf ( Canis lupus ) reintroduction (1998–2005) and evaluated temporal, spatial, and environmental factors that explain variation in adult, subadult, and kitten survival. Using Program MARK and multimodel inference, we modeled cougar survival based on demographic status, season, and landscape attributes. Our best models for adult and independent subadults indicated that females survived better than males and survival increased with age until cougars reached older ages. Lower elevations and increasing density of roads, particularly in areas open to cougar hunting north of Yellowstone National Park (YNP), increased mortality risks for cougars on the GYNR. Indices of ungulate biomass, cougar and wolf population size, winter severity, rainfall, and individual characteristics such as the presence of dependent young, age class, and use of Park or Wilderness were not important predictors of survival. Kitten survival increased with age, was lower during winter, increased with increasing minimum estimates of elk calf biomass, and increased with increasing density of adult male cougars. Using our best model, we mapped adult cougar survival on the GYNR landscape. Results of receiver operating characteristic (ROC) analysis indicated a good model fit for both female (area under the curve [AUC] = 0.81, 95%CI = 0.70–0.92, n  = 35 locations) and male cougars (AUC = 0.84, 95%CI = 0.74–0.94, n  = 49 locations) relative to hunter harvest locations in our study area. Using minimum estimates of survival necessary to sustain the study population, we developed a source-sink surface and we identify several measures that resource management agencies can take to enhance cougar population management based on a source-sink strategy.

Montana, Wyoming

Study design and sampling intensity for demographic analyses of bear populations

The rate of population change through time (??) is a fundamental element of a wildlife population's conservation status, yet estimating it with acceptable precision for bears is difficult. For studies that follow known (usually marked) bears, ?? can be estimated during some defined time by applying either life-table or matrix projection methods to estimates of individual vital rates. Usually however, confidence intervals surrounding the estimate are broader than one would like. Using an estimator suggested by Doak et al. (2005), we explored the precision to be expected in ?? from demographic analyses of typical grizzly (Ursus arctos) and American black (U. americanus) bear data sets. We also evaluated some trade-offs among vital rates in sampling strategies. Confidence intervals around ?? were more sensitive to adding to the duration of a short (e.g., 3 yrs) than a long (e.g., 10 yrs) study, and more sensitive to adding additional bears to studies with small (e.g., 10 adult females/yr) than large (e.g., 30 adult females/yr) sample sizes. Confidence intervals of ?? projected using process-only variance of vital rates were only slightly smaller than those projected using total variances of vital rates. Under sampling constraints typical of most bear studies, it may be more efficient to invest additional resources into monitoring recruitment and juvenile survival rates of females already a part of the study, than to simply increase the sample size of study females. ?? 2011 International Association for Bear Research and Management.

Ursus

Distribution of grizzly bears in the Greater Yellowstone Ecosystem, 2004

The US Fish and Wildlife Service (USFWS) proposed delisting the Yellowstone grizzly bear ( Ursus arctos horribilis ) in November 2005. Part of that process required knowledge of the most current distribution of the species. Here, we update an earlier estimate of occupied range (1990–2000) with data through 2004. We used kernel estimators to develop distribution maps of occupied habitats based on initial sightings of unduplicated females ( n = 481) with cubs of the year, locations of radiomarked bears ( n = 170), and spatially unique locations of conflicts, confrontations, and mortalities ( n = 1,075). Although each data set was constrained by potential sampling bias, together they provided insight into areas in the Greater Yellowstone Ecosystem (GYE) currently occupied by grizzly bears. The current distribution of 37,258 km 2 (1990–2004) extends beyond the distribution map generated with data from 1990–2000 (34,416 km 2 ). Range expansion is particularly evident in parts of the Caribou–Targhee National Forest in Idaho and north of Spanish Peaks on the Gallatin National Forest in Montana.

Wyoming

Changing numbers of spawning cutthroat trout in tributary streams of Yellowstone Lake and estimates of grizzly bears visiting streams from DNA

Spawning Yellowstone cutthroat trout ( Oncorhynchus clarki ) provide a source of highly digestible energy for grizzly bears ( Ursus arctos ) that visit tributary streams to Yellowstone Lake during the spring and early summer. During 1985–87, research documented grizzly bears fishing on 61% of the 124 tributary streams to the lake. Using track measurements, it was estimated that a minimum of 44 grizzly bears fished those streams annually. During 1994, non-native lake trout ( Salvelinus namaycush ) were discovered in Yellowstone Lake. Lake trout are efficient predators and have the potential to reduce the native cutthroat population and negatively impact terrestrial predators that use cutthroat trout as a food resource. In 1997, we began sampling a subset of streams ( n = 25) from areas of Yellowstone Lake surveyed during the previous study to determine if changes in spawner numbers or bear use had occurred. Comparisons of peak numbers and duration suggested a considerable decline between study periods in streams in the West Thumb area of the lake. The apparent decline may be due to predation by lake trout. Indices of bear use also declined on West Thumb area streams. We used DNA from hair collected near spawning streams to estimate the minimum number of bears visiting the vicinity of spawning streams. Seventy-four individual bears were identified from 429 hair samples. The annual number of individuals detected ranged from 15 in 1997 to 33 in 2000. Seventy percent of genotypes identified were represented by more than 1 sample, but only 31% of bears were documented more than 1 year of the study. Sixty-two (84%) bears were only documented in 1 segment of the lake, whereas 12 (16%) were found in 2–3 lake segments. Twenty-seven bears were identified from hair collected at multiple streams. One bear was identified on 6 streams in 2 segments of the lake and during 3 years of the study. We used encounter histories derived from DNA and the Jolly-Seber procedure in Program MARK to produce annual estimates of grizzly bears visiting streams. Approximately 68 grizzly bears visited the vicinity of cutthroat trout spawning streams annually. Thus, approximately 14–21% of grizzly bears in the Greater Yellowstone Ecosystem (GYE) may have used this threatened food resource annually. Yellowstone National Park (YNP) is attempting to control the lake trout population in Yellowstone Lake; our results underscore the importance of that effort to grizzly bears.

Wyoming

Use of naturally occurring mercury to determine the importance of cutthroat trout to Yellowstone grizzly bears

Spawning cutthroat trout ( Oncorhynchus clarki (Richardson, 1836)) are a potentially important food resource for grizzly bears ( Ursus arctos horribilis Ord, 1815) in the Greater Yellowstone Ecosystem. We developed a method to estimate the amount of cutthroat trout ingested by grizzly bears living in the Yellowstone Lake area. The method utilized ( i ) the relatively high, naturally occurring concentration of mercury in Yellowstone Lake cutthroat trout (508 ± 93 ppb) and its virtual absence in all other bear foods ( 6 ppb), ( ii ) hair snares to remotely collect hair from bears visiting spawning cutthroat trout streams between 1997 and 2000, ( iii ) DNA analyses to identify the individual and sex of grizzly bears leaving a hair sample, ( iv ) feeding trials with captive bears to develop relationships between fish and mercury intake and hair mercury concentrations, and ( v ) mercury analyses of hair collected from wild bears to estimate the amount of trout consumed by each bear. Male grizzly bears consumed an average of 5 times more trout/kg bear than did female grizzly bears. Estimated cutthroat trout intake per year by the grizzly bear population was only a small fraction of that estimated by previous investigators, and males consumed 92% of all trout ingested by grizzly bears.

Wyoming, Montana, Idaho

Use of sulfur and nitrogen stable isotopes to determine the importance of whitebark pine nuts to Yellowstone grizzly bears

Whitebark pine ( Pinus albicaulis ) is a masting species that produces relatively large, fat- and protein-rich nuts that are consumed by grizzly bears ( Ursus arctos horribilis ). Trees produce abundant nut crops in some years and poor crops in other years. Grizzly bear survival in the Greater Yellowstone Ecosystem is strongly linked to variation in pine-nut availability. Because whitebark pine trees are infected with blister rust ( Cronartium ribicola ), an exotic fungus that has killed the species throughout much of its range in the northern Rocky Mountains, we used stable isotopes to quantify the importance of this food resource to Yellowstone grizzly bears while healthy populations of the trees still exist. Whitebark pine nuts have a sulfur-isotope signature (9.2 ± 1.3‰ (mean ± 1 SD)) that is distinctly different from those of all other grizzly bear foods (ranging from 1.9 ± 1.7‰ for all other plants to 3.1 ± 2.6‰ for ungulates). Feeding trials with captive grizzly bears were used to develop relationships between dietary sulfur-, carbon-, and nitrogen-isotope signatures and those of bear plasma. The sulfur and nitrogen relationships were used to estimate the importance of pine nuts to free-ranging grizzly bears from blood and hair samples collected between 1994 and 2001. During years of poor pine-nut availability, 72% of the bears made minimal use of pine nuts. During years of abundant cone availability, 8 ± 10% of the bears made minimal use of pine nuts, while 67 ± 19% derived over 51% of their assimilated sulfur and nitrogen (i.e., protein) from pine nuts. Pine nuts and meat are two critically important food resources for Yellowstone grizzly bears.

Wyoming

Estimating total human-caused mortality from reported mortality using data from radio-instrumented grizzly bears

Tracking mortality of the Yellowstone grizzly bear ( Ursus arctos horribilis ) is an essential issue of the recovery process. Problem bears removed by agencies are well documented. Deaths of radiocollared bears are known or, in many cases, can be reliably inferred. Additionally, the public reports an unknown proportion of deaths of uncollared bears. Estimating the number of non-agency human-caused mortalities is a necessary element that must be factored into the total annual mortality. Here, we describe a method of estimating the number of such deaths from records of reported human-caused bear mortalities. We used a hierarchical Bayesian model with a non-informative prior distribution for the number of deaths. Estimates of reporting rates developed from deaths of radio-instrumented bears from 1983 to 2000 were used to develop beta prior probability distributions that the public will report a death. Twenty-seven known deaths of radio-instrumented bears occurred during this period with 16 reported. Additionally, fates of 23 radio-instrumented bears were unknown and are considered possible unreported mortalities. We describe 3 ways of using this information to specify prior distributions on the probability a death will be reported by the public. We estimated total deaths of noninstrumented bears in running 3-year periods from 1993 to 2000. Thirty-nine known deaths of non-instrumented bears were reported during this period, ranging from 0 to 7/year. Seven possible mortalities were recorded. We applied the method to both sets of mortality data. Results from this method can be combined with agency removals and deaths of collared bears to produce defensible estimates of total mortality over relevant periods and to incorporate uncertainty when evaluating mortality limits established for the Yellowstone grizzly bear population. Assumptions and limitations of this procedure are discussed.

Idaho, Montana, Wyoming