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Double-survey estimates of bald eagle populations in Oregon

The literature on abundance of birds of prey is almost devoid of population estimates with statistical rigor. Therefore, we surveyed bald eagle (Haliaeetus leucocephalus) populations on the Crooked and lower Columbia rivers of Oregon and used the double-survey method to estimate populations and sighting probabilities for different survey methods (aerial, boat, vehicle) and bald eagle ages (adults vs. subadults). Sighting probabilities were consistently <1.00, so direct counts were biased estimates of abundance. There was a greater probability of observing adult (0.625) versus subadult (0.494) eagles on the Crooked River for different survey methods, but we did not detect differences in sighting probabilities for adults (0.707) versus subadults (0.683) on the Columbia River. The probability of observing bald eagles from aircraft (0.636) was lower than that from boats (0.758) on the Columbia River, and sighting probabilities from aircraft (0.354) were lower than those from the ground (0.764) on the Crooked River. Precision of population estimates was generally good: most (68%, n = 30) coefficients of variation were <10%, and only 7% were >20%. The results revealed variable and negative bias (percent relative bias = -9 to -70%) of direct counts and emphasized the importance of estimating populations where some measure of precision and ability to conduct inference tests are available. We recommend use of the double-survey method to estimate abundance of bald eagle populations and other raptors in open habitats.

Journal of Wildlife Management↗

Effects of saline drinking water on early gosling development

Relatively high levels of saline drinking water may adversely affect the growth, development, and survival of young waterfowl. Saline drinking water was suspect in the low survival rate of Canada goose (Branta canadensis) goslings at Fish Springs National Wildlife Refuge (FSNWR) in western Utah. Hence, we investigated the effects of saline drinking water on the survival and growth of captive, wild-strain goslings from day 1-28 following hatch. We compared survival and growth (as measured by body mass, wing length, and culmen length) between a control group on tap water with a mean specific conductivity of 650 μS/cm, and 2 saline water treatments: (1) intermediate level (12,000 μS/cm), and (2) high level (18,000 μS/cm). Gosling mortality occurred only in the 18,000 μS/cm treatment group (33%; n = 9). Slopes of regressions of mean body mass, wing length, and culmen length on age were different from each other (P < 0.05), except for culmen length for the intermediate and high treatment levels. We predict that free-ranging wild goslings will experience mortality at even lower salinity levels than captive goslings because of the combined effects of depressed growth and environmental stresses, including hot desert temperatures and variable food quality over summer.

Journal of Wildlife Management↗

Autocorrelation of location estimates and the analysis of radiotracking data

The wildlife literature has been contradictory about the importance of autocorrelation in radiotracking data used for home range estimation and hypothesis tests of habitat selection. By definition, the concept of a home range involves autocorrelated movements, but estimates or hypothesis tests based on sampling designs that predefine a time frame of interest, and that generate representative samples of an animal's movement during this time frame, should not be affected by length of the sampling interval and autocorrelation. Intensive sampling of the individual's home range and habitat use during the time frame of the study leads to improved estimates for the individual, but use of location estimates as the sample unit to compare across animals is pseudoreplication. We therefore recommend against use of habitat selection analysis techniques that use locations instead of individuals as the sample unit. We offer a general outline for sampling designs for radiotracking studies.

Journal of Wildlife Management↗

Effects of egg and hatchling harvest on American alligators in Florida

Harvest of crocodilian eggs and young for captive rearing (ranching) has been used worldwide as an option for producing crocodilian skins and meat from wild stock. The long-term effects of harvesting a certain proportion of early age class, wild American alligators (Alligator mississippiensis) without repatriation is unknown. We removed an estimated 50% of annual production of alligators on Lakes Griffin and Jesup in central Florida over an 11-year period and monitored population levels via night-light counts. Densities of the total alligator population increased (P < 0.037) on all areas. Count densities of adult (≥183 cm total length [TL]) alligators increased (P < 0.003) on harvest areas but remained stable (P = 0.830) on the control (no harvest) area, Lake Woodruff National Wildlife Refuge (Lake Woodruff NWR). Observed densities of juvenile (<122 cm TL) alligators remained stable (P > 0.117), and subadult (122-182 cm TL) alligators increased (P < 0.011) on harvest areas. The density of juveniles on the control area increased (P = 0.006), and the density of subadults showed some evidence of increasing (P = 0.088). No changes were detected in size distributions on the treatment areas. Nest production, as observed from aerial helicopter surveys, increased (P < 0.039) on Lake Woodruff NWR and Lake Jesup and showed some evidence of an increase on Lake Griffin (P = 0.098) during 1983-91. A 50% harvest rate of eggs or hatchlings did not adversely affect recruitment into the subadult or adult size classes.

Journal of Wildlife Management↗

Activity patterns and time budgets of the declining sea otter population at Amchitka Island, Alaska

Time budgets of predators may reflect population status if time spent foraging varies with local prey abun- dance. We assumed that the sea otter (Enhydra lutris) population at Amchitka Island, Alaska, USA, had been at equilibrium since the early 1960s and collected time budgets of otters to be used to represent future conditions of currently expanding sea otter populations. We used radiotelemetry to monitor activity-time budgets of otters from August 1992 to March 1994. Sea otter activity was directly linked to sex, age, weather condition, season, and time of day. Sea otters differed in percent time foraging among cohorts but not within cohorts. Percent time foraging ranged from 21% for females with very young (≤ 3weeks of age) dependent pups to 52% for females with old (≥10 weeks of age) pups. Otters foraged more and hauled out more as local sea conditions worsened. Adult males spent less time foraging during winter and spring, consistent with seasonal changes in prey selection. Time spent for- aging was similar to that reported for otters in California and an established population in Prince William Sound, Alaska, but greater than that of otters in recently established populations in Oregon and Alaska. Despite current evidence indicating that the population was in decline during our study, we were unable to recognize this change using time budgets. Our results illustrate the importance of stratifying analyses of activity patterns by age and sex cohorts and the complexity inherent in comparisons of behavioral data between different populations relying on distinct prey bases.

Journal of Wildlife Management↗

The northern Yellowstone elk: density dependence and climatic conditions

We analyzed a time series of estimates of elk ( Cervus elaphus ) numbers on the northern Yellowstone winter range from 1964 to 1979 and 1986 to 1995 using a variety of discrete time stochastic population dynamic models. These models included adjustments for density, an increase in the area of winter range used by elk, lagged effects of the weather covariates of spring precipitation, snow depth and winter temperature, and the impacts of the 1988 drought and fires. An information-criteria-based model-selection process strongly supported evidence of density dependence. The best model, a Ricker model, distinguished between the 2 time periods. The bulk of the difference between the 2 periods is attributed to an increase in the amount of winter range used by elk. Inclusion of the covariates spring precipitation and spring precipitation squared greatly improved the model fit. We detected a short-lived increase in elk population growth rate following the 1988 drought and fires. Fertility and survivorship of adults appeared to have different density-dependent forms that together result in a biphasic relationships between population growth rate and density. This study confirms the presence of density-dependent regulation in the norther Yellowstone elk herd, and enhances our understanding of population dynamics of these ungulates.

Journal of Wildlife Management↗

Distribution and movements of female northern pintails radiotagged in San Joaquin Valley, California

To improve understanding of northern pintail (Anas acuta) distribution in central California (CCA), we radiotagged 191 Hatch-Year (HY) and 228 After-Hatch-Year (AHY) female northern pintails during late August-early October, 1991-1993, in the San Joaquin Valley (SJV) and studied their movements through March each year. Nearly all (94.3%) wintered in CCA, but 5.7% went to southern California, Mexico, or unknown areas; all that went south left before hunting season. Of the 395 radiotagged pintails that wintered in CCA, 83% flew from the SJV north to other CCA areas (i.e., Sacramento Valley [SACV], Sacramento-San Joaquin River Delta [Delta], Suisun Marsh, San Francisco Bay) during September-January; most went during December. Movements coincid- ed with start of hunting seasons and were related to pintail age, mass, capture location, study year, and weather. Among pintails with less than average mass, AHY individuals tended to leave the SJV earlier than HY individuals. Weekly distribution was similar among capture locations and years but a greater percentage of pintails radiotagged in Tulare Basin (south part of SJV) were known to have (10.3% vs. 0.9%) or probably (13.8% vs. 4.6%) wintered south of CCA than pintails radiotagged in northern SJV areas (i.e., Grassland Ecological Area [EA] and Mendota Wildlife Area [WA]). Also, a greater percentage of SJV pintails went to other CCA areas before hunting season in the drought year of 1991-1992 than later years (10% vs. 3-5%). The percent of radiotagged pintails from Grass- land EA known to have gone south of CCA also was greater during 1991-1992 than later years (2% vs. 0%), but both the known (19% vs. 4%) and probable (23% vs. 12%) percent from Tulare Basin that went south was greatest during 1993-1994, when availability of flooded fields there was lowest. The probability of pintails leaving the SJV was 57% (95% CI = 8-127%) greater on days with than without rain, and more movements per bird out of SJV occurred in years with more rain and fog but fewer days with southerly winds. Movements by pintails and changes in pintail distributions, direct recovery distributions, and harvest rates suggest the disproportionate decline of pin- tails in Tulare Basin was due to a lower percentage of pintails moving there in fall and a greater percentage or ear- lier movements north and south out of Tulare Basin. With fewer in Tulare Basin to replace Grasslands EA pintails going north in December, pintail abundance in the northern SJV declined during late winter. Changes in move- ment patterns correspond to habitat loss in Tulare Basin and increased habitats in SACV and western mainland Mexico. Habitat improvements, especially in Tulare Basin, that increase food, sanctuary, and winter survival would probably help restore pintails throughout the SJV.

Journal of Wildlife Management↗

Estimation of brood and nest survival: Comparative methods in the presence of heterogeneity

The Mayfield method has been widely used for estimating survival of nests and young animals, especially when data are collected at irregular observation intervals. However, this method assumes survival is constant throughout the study period, which often ignores biologically relevant variation and may lead to biased survival estimates. We examined the bias and accuracy of 1 modification to the Mayfield method that allows for temporal variation in survival, and we developed and similarly tested 2 additional methods. One of these 2 new methods is simply an iterative extension of Klett and Johnson's method, which we refer to as the Iterative Mayfield method and bears similarity to Kaplan-Meier methods. The other method uses maximum likelihood techniques for estimation and is best applied to survival of animals in groups or families, rather than as independent individuals. We also examined how robust these estimators are to heterogeneity in the data, which can arise from such sources as dependent survival probabilities among siblings, inherent differences among families, and adoption. Testing of estimator performance with respect to bias, accuracy, and heterogeneity was done using simulations that mimicked a study of survival of emperor goose ( Chen canagica ) goslings. Assuming constant survival for inappropriately long periods of time or use of Klett and Johnson's methods resulted in large bias or poor accuracy (often >5% bias or root mean square error) compared to our Iterative Mayfield or maximum likelihood methods. Overall, estimator performance was slightly better with our Iterative Mayfield than our maximum likelihood method, but the maximum likelihood method provides a more rigorous framework for testing covariates and explicity models a heterogeneity factor. We demonstrated use of all estimators with data from emperor goose goslings. We advocate that future studies use the new methods outlined here rather than the traditional Mayfield method or its previous modifications.

Journal of Wildlife Management↗

Survival of Columbian white-tailed deer in western Oregon

Columbian white-tailed deer (Odocoileus virginianus leucurus; CWTD) are an endangered subspecies on which little demographic information exists. We determined survival rates and causes of mortality for 64 radiocol- lared adults from 1996 to 1998, and for 63 radiocollared neonatal fawns during the summer and fall months of 1996-2001 in Douglas County, Oregon, USA. Annual adult survival rates averaged 0.74 over 3 years, and most mor- tality (73%) occurred between fall and winter. Seasonal survival was lowest (0.75) for the fall-winter 1997-1998, and was 20.90 during all spring-summer periods. Annual and seasonal survival rates did not differ by gender. Average annual survival was 0.77 for deer in wildland areas compared with 0.66 for deer in suburban areas, but these dif- ferences were not consistent between years and seasons. Survival over the entire 3-year study was low (0.38). Eight deer died from a combination of emaciation and disease, and almost all (92%) necropsied deer were in poor body condition. Fawn survival to 7 months was low (0.14, 95% CI = 0.02-0.26) and declined most rapidly during the first 1.5 months of life. Predation (n = 21) and abandonment (n = 6) were the most frequent known causes of death for fawns. Our results suggest that CWTD may have responded to density-dependent factors during this short-term study, although the effects of other environmental or intrinsic factors cannot be ignored. Fawn survival may be insufficient to produce enough recruits for population growth and eventual range expansion.

Journal of Wildlife Management↗

Identifying predators and fates of grassland passerine nests using miniature video cameras

Nest fates, causes of nest failure, and identities of nest predators are difficult to determine for grassland passerines. We developed a miniature video-camera system for use in grasslands and deployed it at 69 nests of 10 passerine species in North Dakota during 1996-97. Abandonment rates were higher at nests <1 day after camera deployment (23%) than <1 day after nests were found (2%, P = 0.001). Most birds returned to nests 30 min after camera deployment, but clay-colored sparrows (Spizella pallida) took longer to return to nests than other species (P = 0.035). Cameras did not appear to increase nest predation rates. We videotaped predation by 11 species at 29 nests: eggs or nestlings were destroyed by mice, ground squirrels, weasel, badgers, canids, deer, cowbirds, and hawks. All eggs or nestlings were removed in <15 min at 14 depredated nests. Contents were removed during >1 day or night (22-116 hr) at 6 nests, 5 of which were depredated by ground squirrels or mice. For nests without cameras, estimated predation rates were lower for ground nests than aboveground nests (P = 0.055), but did not differ between open and covered nests (P = 0.74). Open and covered nests differed, however, when predation risk (estimated by initial-predation rate) was examined separately for day and night using camera-monitored nests; the frequency of initial predations that occurred during the day was higher for open nests than covered nests (P = 0.015). Thus, vulnerability of some nest types may depend on the relative importance of nocturnal and diurnal predators. Predation risk increased with nestling age from 0 to 8 days (P = 0.07). Up to 15% of fates assigned to camera-monitored nests were wrong when based solely on evidence that would have been available from periodic nest visits. There was no evidence of disturbance at nearly half the depredated nests, including all 5 depredated by large mammals. Overlap in types of sign left by different predator species, and variability of sign within species, suggests that evidence at nests is unreliable for identifying predators of grassland passerines.

Journal of Wildlife Management↗

Do wolves affect white-tailed buck harvest in northeastern Minnesota?

We used simple linear regression to analyze 8-23 years of data on a wolf (Canis lupus) population and human harvest of white-tailed deer (Odocoileus virginianus) bucks in northeastern Minnesota to determine any effects of wolves on buck harvesting. Over the long term, wolves accounted for at least 14-22% of the inter-year variation in buck harvest in the region, but an unknown amount of variation in hunter effort may have obscured any more precise estimate. For part of the area with poorest habitat, we found strong inverse relationships (r2 = 0.66-0.84) between annual wolf numbers and buck harvests from 1988 to 1995 when hunting pressure was considered relatively constant. However, in better habitat, where our buck harvest sample was larger, we found no evidence of wolves influencing buck harvest. Our findings tend to confirm the suitability of the Minnesota Department of Natural Resource's deer harvest regulations for a sustainable yield.

Journal of Wildlife Management↗

Effect of hibernation and reproductive status on body mass and condition of coastal brown bears

We investigated the effect of hibernation and reproductive status on changes in body mass and composition of adult female brown bears (Ursus arctos) on the Kenai Peninsula, Alaska. This information is fundamental to understanding nutritional ecology of wild brown bear populations. Six adult females handled in the fall and following spring (paired samples) lost 73 &plusmn; 22 kg (x̄ &plusmn; SD; 32 &plusmn; 10%) of fall body mass over 208 &plusmn; 19 days. Of this mass loss, 56 &plusmn; 22% (55 &plusmn; 22 kg) was lipid and 44 &plusmn; 22% (43 &plusmn; 21 kg) was lean body mass. Catabolism of lipid stores accounted for 88.4 &plusmn; 8.1% of the body energy used to meet maintenance demands. Overwinter differences in body composition of adult females assessed only once in either the fall (n = 21) or spring (n = 32) were similar to those of paired samples. Relative fatness of bears entering the den was positively related to the contribution of fat (%) to body mass (P < 0.01) and body energy (P < 0.01) losses during hibernation. Thus, relative fatness at the onset of fasting influences the relative proportion of lipid stores and lean body mass catabolized to meet protein and energy demands during hibernation. In the spring, lone females had greater body and lean masses than females with cubs of the year or yearlings. Lipid content was greatest in lone females in the fall. Studies using body mass and composition as indices of population health should consider season or reproductive class.

Journal of Wildlife Management↗

Survival of American black ducks radiomarked in Quebec, Nova Scotia, and Vermont

We monitored survival of 397 radiomarked juvenile American black ducks (Anas rubripes) distributed among Les Escoumins ( n = 75) and Kamouraska, Quebec ( n = 84), Amherst Point, Nova Scotia ( n = 89), and a site on the Vermont-Quebec border ( n = 149) during autumn 1990 and 1991. Eighty-six percent (215 of 250) of all confirmed mortalities during the study was from hunting; 72% of marked ducks were shot and retrieved and 14% were shot and unretrieved. We tested for differences in survival in relation to sex, body mass, year (1990-91, 1991-92), and among the 4 locations for each of 2 monitoring periods (early, EMP; late, LMP). With data from the EMP for Vermont-Quebec in 1990 and 1991, Les Escoumins in 1990, and Amherst Point in 1991, survival of hatching-year (HY) males and females did not differ ( P = 0.357). For sexes combined for the EMP, survival of ducks was greater in 1991 than 1990 ( P = 0.086), and differed among locations ( P = 0.013). Survival (years combined) was greater at Amherst Point than at Kamouraska ( P = 0.003) and Vermont-Quebec ( P = 0.002) during the EMP. The highest survival rate at Amherst Point (0.545 ± 0.056 [SE]) was associated with the latest date (8 Oct) of season opening; the lowest survival rate (0.395 ± 0.043) was at the Vermont-Quebec border, where hunter numbers and activity were greatest. For the LMP, no interaction between years and locations was detected ( P = 0.942), and no differences in survival existed between years ( P = 0.102) and among locations ( P = 0.349). No association was detected between body mass at capture and survival of combined males and females during the EMP ( P = 0.572) or during the LMP ( P = 0.965). When we censored hunting losses for combined years for each period, EMP or LMP, all survival estimates exceeded 0.800 (0.809-0.965). These data emphasize need for an improved harvest strategy for American black ducks in North America to allow for increases in breeding populations to achieve population goals.

Nova Scotia, Quebec, Vermont↗

Annual survival and site fidelity of Stellar's Eiders molting along the Alaska Peninsula

Populations of Steller's eiders ( Polysticta stelleri ) molting and wintering along the Alaska Peninsula have declined since the 1960's. We captured and marked a large sample of Steller's eiders molting in 2 lagoons along the Alaska Peninsula between 1975-97. We used mark-recapture analysis techniques to estimate annual survival and movement probabilities within and among lagoons for male and female eiders. Estimates of annual survival (±SE) were 0.899 ± 0.032 for females and 0.765 ± 0.044 for males. Both sexes showed high rates of fidelity to specific molting locations (>95%) within lagoons; yet we found no evidence that annual probability of survival differed among groups molting in different locations either within or among lagoons. We found weak evidence that annual survival decreased between the periods 1975-81 and 1991-97. The lower survival of males compared to females is unusual for waterfowl and may result in a female-biased sex ratio. We conclude that a decrease in adult survival may have initiated the long-term population decline. Further, a shortage of males may be limiting reproductive potential.

Alaska↗

Simultaneous use of mark-recapture and radiotelemetry to estimate survival, movement, and capture rates

Biologists often estimate separate survival and movement rates from radio-telemetry and mark-recapture data from the same study population. We describe a method for combining these data types in a single model to obtain joint, potentially less biased estimates of survival and movement that use all available data. We furnish an example using wood thrushes ( Hylocichla mustelina ) captured at the Piedmont National Wildlife Refuge in central Georgia in 1996. The model structure allows estimation of survival and capture probabilities, as well as estimation of movements away from and into the study area. In addition, the model structure provides many possibilities for hypothesis testing. Using the combined model structure, we estimated that wood thrush weekly survival was 0.989 ± 0.007 ( ± SE). Survival rates of banded and radio-marked individuals were not different ( α [ S radioed , S banded ] =log[ S radioed / S banded ]=0.0239 ± 0.0435). Fidelity rates (weekly probability of remaining in a stratum) did not differ between geographic strata ( Ψ = 0.911 ± 0.020; α [ Ψ 11 , Ψ 22 ]=0.0161 ± 0.047), and recapture rates ( p = 0.097 ± 0.016) banded and radio-marked individuals were not different ( α [ P radioed , P banded ]=0.145 ± 0.655). Combining these data types in a common model resulted in more precise estimates of movement and recapture rates than separate estimation, but ability to detect stratum or mark-specific differences in parameters was week. We conducted simulation trials to investigate the effects of varying study designs on parameter accuracy and statistical power to detect important differences. Parameter accuracy was high (relative bias [RBIAS] <2 %) and confidence interval coverage close to nominal, except for survival estimates of banded birds for the 'off study area' stratum, which were negatively biased (RBIAS -7 to -15%) when sample sizes were small (5-10 banded or radioed animals 'released' per time interval). To provide adequate data for useful inference from this model, study designs should seek a minimum of 25 animals of each marking type observed (marked or observed via telemetry) in each time period and geographic stratum.

Journal of Wildlife Management↗

Declining scaup populations: A retrospective analysis of long-term population and harvest survey data

We examined long-term databases concerning population status of scaup (lesser [ Aythya affinis ] and greater scaup [ A. marila ] combined) and harvest statistics of lesser scaup to identify factors potentially limiting population growth. Specifically, we explored evidence for and against the general hypotheses that scaup populations have declined in association with declining recruitment and/or female survival. We examined geographic heterogeneity in scaup demographic patterns that could yield evidence about potential limiting factors. Several biases exist in survey methodology used to estimate scaup populations and harvest statistics; however, none of these biases likely accounted for our major findings that (1) the continental scaup breeding population has declined over the last 20 years, with widespread and consistent declines within surveyed areas of the Canadian western boreal forest where most lesser scaup breed; (2) sex ratios of lesser scaup in the U.S. harvest have increased (more males now relative to females); and (3) age ratios of lesser scaup in the U.S. harvest have declined (fewer immatures now relative to adults), especially in the midcontinent region. We interpreted these major findings as evidence that (1) recruitment of lesser scaup has declined over the last 20 years, particularly in the Canadian western boreal forest; and (2) survival of female lesser scaup has declined relative to that of males. We found little evidence that harvest was associated with the scaup population decline. Our findings underscore the need for both improvements and changes to population survey procedures and new research to discriminate among various hypotheses explaining the recent scaup population decline.

North America↗

Winter severity and wolf predation on a formerly wolf-free elk herd

We studied wolf (Canis lupus) predation on elk (Cervus elaphus) in Yellowstone National Park from 17 March to 15 April 1997 (severe winter conditions) and from 2 to 31 March 1998 (mild winter conditions) 2-3 years after wolves were reintroduced to the park. Elk composed 91 % of 117 kills. Data comparisons for 1997 versus 1998 were: hunting success rate, 26% versus 15%; kill rate, 17.1 kg/wolf/day versus 6.1; percent of kill consumed in first day, 7 versus 86; percent femur marrow fat of adult kills, 27 versus 70; calf:adult ratios of kills, 2:33 versus 17:23; sex ratio of kills, 14M:19F versus 17M:6F; mean age of elk killed, males 6.1 years, females 15.2 versus males, 4.8, females 13.0. Winter severity influenced the wolf-elk relationship more than the naivete of the elk herd to predation by wolves.

Journal of Wildlife Management↗