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Charles C. Schwartz

Publications and source records attributed to Charles C. Schwartz.

30 records · Page 2Linked to original sources

Nutritional ecology of ursids: A review of newer methods and management implications

The capability to understand the nutritional ecology of free-ranging bears has increased dramatically in the last 20 years. Advancements have occurred because (1) managers and biologists recognized the need to link habitat quality, productivity, and variability with bear movements, home ranges, and demographic parameters like reproductive output, survival, and population growth, and (2) several research teams are using new methods to build on the results of earlier field studies. Our ability to couple new field methods and empirical field research with controlled experiments using captive bears has been central to our increased understanding of bear nutrition. Newer methods include the use of stable isotopes to quantify assimilated diet and nutrient flows within ecosystems, bioelectrical impedance to measure body composition, and naturally occurring mercury to estimate fish intake. Controlled experiments using captive bears have been integral to developing methods, isolating specific variables by controlling the environment, and providing additional nutritional understanding necessary to interpret field observations. We review new methods and apply our increased understanding of bear nutritional ecology to 3 management issues: (1) the importance of salmon ( Oncorhynchus spp.) to brown bears ( Ursus arctos ) in the Pacific Northwest, (2) the consequences of the closure of the Yellowstone garbage dumps to grizzly bears, and (3) the relocation of problem bears.

Ursus

Modeling survival: application of the Andersen-Gill model to Yellowstone grizzly bears

Wildlife ecologists often use the Kaplan-Meier procedure or Cox proportional hazards model to estimate survival rates, distributions, and magnitude of risk factors. The Andersen-Gill formulation (A-G) of the Cox proportional hazards model has seen limited application to mark-resight data but has a number of advantages, including the ability to accommodate left-censored data, time-varying covariates, multiple events, and discontinuous intervals of risks. We introduce the A-G model including structure of data, interpretation of results, and assessment of assumptions. We then apply the model to 22 years of radiotelemetry data for grizzly bears ( Ursus arctos ) of the Greater Yellowstone Grizzly Bear Recovery Zone in Montana, Idaho, and Wyoming, USA. We used Akaike's Information Criterion (AIC c ) and multi-model inference to assess a number of potentially useful predictive models relative to explanatory covariates for demography, human disturbance, and habitat. Using the most parsimonious models, we generated risk ratios, hypothetical survival curves, and a map of the spatial distribution of high-risk areas across the recovery zone. Our results were in agreement with past studies of mortality factors for Yellowstone grizzly bears. Holding other covariates constant, mortality was highest for bears that were subjected to repeated management actions and inhabited areas with high road densities outside Yellowstone National Park. Hazard models developed with covariates descriptive of foraging habitats were not the most parsimonious, but they suggested that high-elevation areas offered lower risks of mortality when compared to agricultural areas.

Journal of Wildlife Management

Mitochondrial phylogeography of moose (Alces alces) in North America

Nucleotide variation was assessed from the mitochondrial control region of North American moose ( Alces alces ) to test predictions of a model of range expansion by stepping-stone dispersal and to determine whether patterns of genetic variation support the current recognition of 4 subspecies. Haplotypes formed a star phylogeny indicative of a recent expansion of populations. Values of nucleotide and haplotype diversity were low continentwide but were greatest in the central part of the continent and lowest in peripheral populations. Despite low mitochondrial diversity, moose exhibited a high degree of differentiation regionally, which was not explained by isolation by distance. Our data indicate a pattern of colonization consistent with a large central population that supplied founders to peripheral populations (other than Alaska), perhaps through rare, long-distance dispersal events (leptokurtic dispersal) rather than mass dispersal by a stepping-stone model. The colonization scenario does not account for the low haplotype diversity observed in Alaska, which may be derived from a postcolonization bottleneck. Establishment of peripheral populations by leptokurtic dispersal and subsequent local adaptation may have been sufficient for development of morphological differentiation among extant subspecies.

Journal of Mammalogy

Large carnivores, moose, and humans: A changing paradigm of predator management in the 21st century

We compare and contrast the evolution of human attitudes toward large carnivores between Europe and North America. In general, persecution of large carnivores began much earlier in Europe than North America. Likewise, conservation programs directed at restoration and recovery appeared in European history well before they did in North America. Together, the pattern suggests there has been an evolution in how humans perceive large predators. Our early ancestors were physically vulnerable to large carnivores and developed corresponding attitudes of respect, avoidance, and acceptance. As civilization evolved and man developed weapons, the balance shifted. Early civilizations, in particular those with pastoral ways, attempted to eliminate large carnivores as threats to life and property. Brown bears (Ursus arctos) and wolves (Canis lupus) were consequently extirpated from much of their range in Europe and in North America south of Canada. Efforts to protect brown bears began in the late 1880s in some European countries and population reintroductions and augmentations are ongoing. They are less controversial than in North America. On the other hand, there are no wolf introductions, as has occurred in North America, and Europeans have a more negative attitude towards wolves. Control of predators to enhance ungulate harvest varies. In Western Europe, landowners own the hunting rights to ungulates. In the formerly communistic Eastern European countries and North America, hunting rights are held in common, although this is changing in some Eastern European countries. Wolf control to increase harvests of moose (Alces alces) occurs in parts of North America and Russia; bear control for similar reasons only occurs in parts of North America. Surprisingly, bears and wolves are not controlled to increase ungulates where private landowners have the hunting rights in Europe, although wolves were originally exterminated from these areas. Both the inability of scientific research to adequately predict the effect of predator control on ungulate populations and a shift in public attitudes toward large carnivores have resulted in an accelerating number of challenges to predator management in places where it is still espoused. Utilitarian attitudes towards wildlife are declining in Western cultures and people now increasingly recognize the intrinsic value of wildlife, including large predators. In the future, agencies responsible for managing resident wildlife will face increased pressure to balance the needs of the hunting public with the desires of non-hunting publics. We suggest that in the next century we will witness a continued shift in how wildlife agencies manage both moose and large carnivores. More attention will be paid to maintaining and restoring intact ecosystems and less toward sustainable yield of meat.

Alces

Reproductive maturation and senescence in the female brown bear

Changes in age-specific reproductive rates can have important implications for managing populations, but the number of female brown (grizzly) bears ( Ursus arctos ) observed in any one study is usually inadequate to quantify such patterns, especially for older females and in hunted areas. We examined patterns of reproductive maturation and senescence in female brown bears by combining data from 20 study areas from Sweden, Alaska, Canada, and the continental United States. We assessed reproductive performance based on 4,726 radiocollared years for free-ranging female brown bears (age ≥3); 482 of these were for bears ≥20 years of age. We modeled age-specific probability of litter production using extreme value distributions to describe probabilities for young- and old-age classes, and a power distribution function to describe probabilities for prime-aged animals. We then fit 4 models to pooled observations from our 20 study areas. We used Akaike's Information Criterion (AIC) to select the best model. Inflection points suggest that major shifts in litter production occur at 4-5 and 28-29 years of age. The estimated model asymptote (0.332, 95% CI = 0.319-0.344) was consistent with the expected reproductive cycle of a cub litter every 3 years (0.333). We discuss assumptions and biases in data collection relative to the shape of the model curve. Our results conform to senescence theory and suggest that female age structure in contemporary brown bear populations is considerably younger than would be expected in the absence of modern man. This implies that selective pressures today differ from those that influenced brown bear evolution.

Ursus

Mitochondrial phylogeography of moose ( Alces alces ): Late Pleistocene divergence and population expansion

We examined phylogeographic relationships of moose ( Alces alces ) worldwide to test the proposed existence of two geographic races and to infer the timing and extent of demographic processes underpinning the expansion of this species across the Northern Hemisphere in the late Pleistocene. Sequence variation within the left hypervariable domain of the control region occurred at low or moderate levels worldwide and was structured geographically. Partitioning of genetic variance among regions indicated that isolation by distance was the primary agent for differentiation of moose populations but does not support the existence of distinct eastern and western races. Levels of genetic variation and structure of phylogenetic trees identify Asia as the origin of all extant mitochondrial lineages. A recent coalescence is indicated, with the most recent common ancestor dating to the last ice age. Moose have undergone two episodes of population expansion, likely corresponding to the final interstade of the most recent ice age and the onset of the current interglacial. Timing of expansion for the population in the Yakutia–Manchuria region of eastern Asia indicates that it is one of the oldest populations of moose and may represent the source of founders of extant populations in North America, which were colonized within the last 15,000 years. Our data suggest an extended period of low population size or a severe bottleneck prior to the divergence and expansion of extant lineages and a recent, less-severe bottleneck among European lineages. Climate change during the last ice age, acting through contraction and expansion of moose habitat and the flooding of the Bering land bridge, undoubtedly was a key factor influencing the divergence and expansion of moose populations.

Alaska, Colorado

Distribution of grizzly bears in the Greater Yellowstone Ecosystem, 1990-2000

The Yellowstone grizzly bear ( Ursus arctos horribilis ) has been expanding its range during the past 2 decades and now occupies historic habitats that had been vacant. A current understanding of the distribution of grizzly bears within the ecosystem is useful in the recovery process and to help guide the state and federal land management agencies and state wildlife agencies of Idaho, Wyoming, and Montana as they prepare management plans. We used kernel estimators to develop distribution maps of occupied habitats based on initial sightings of unduplicated females (n = 300) with cubs-of-the-year, information from radiomarked bears (n = 105), and locations of conflicts, confrontations, and mortalities (n = 1,235). Although each data set was constrained by potential sampling bias, together they provided insight into areas within the Greater Yellowstone Ecosystem (GYE) currently occupied by grizzly bears. The current distribution (1990-2000) extends beyond the recovery zone identified in the U.S. Fish and Wildlife Service (USFWS) Recovery Plan. Range expansion is particularly evident in the southern portion of the ecosystem in Wyoming. A comparison of our results from the 1990s to previously published distribution maps show an approximate increase in occupied habitat of 48% and 34% from the 1970s and 1980s, respectively. We discuss data biases and problems implicit to the analysis.

Wyoming

Grizzly bear denning and potential conflict areas in the Greater Yellowstone Ecosystem

Increasing winter use of steep, high-elevation terrain by backcountry recreationists has elevated concern about disturbance of denning grizzly bears (Ursus arctos) in the Greater Yellowstone Ecosystem (GYE). To help identify areas where such conflicts might occur, we developed a spatially explicit model to predict potential denning areas in the GYE. Using a scan area of 630 m around each location, we assigned site attributes to 344 den locations of radio-trackedg rizzly bears from 1975-99. Attributesi dentified as predictorsf or the analysis included elevation, slope, an index of solar radiation, and forest cover. We used the Mahalanobis distance statistic to model the similarity between sites used by denning bears and each cell in the data layers. We used the final Mahalanobis distance model to produce maps of the study area. Potential denning habitat, based upon the model, is abundantw ithin the GYE. Ourr esultsc an be used by land managementa gencies to identifyp otentialc onflict sites and minimize effects of regulated activities on denning grizzly bears. We illustrate how the Gallatin National Forest (GNF) used the model to examine the overlap between potential snowmobile use areas and potential denning habitat as part of a Biological Assessment submitted to the U.S. Fish and Wildlife Service.

Wyoming

Grizzly bear denning chronology and movements in the Greater Yellowstone Ecosystem

Den entrance and emergence dates of grizzly bears ( Ursus arctos ) in the Greater Yellowstone Ecosystem are important to management agencies that wish to minimize impacts of human activities on bears. Current estimates for grizzly bear denning events use data that were collected from 1975–80. We update these estimates by including data obtained from 1981–99. We used aerial telemetry data to estimate week of den entry and emergence by determining the midpoint between the last known active date and the first known date denned, as well as the last known date denned and the first known active date. We also investigated post emergence movement patterns relative to den locations. Mean earliest and latest week of den entry and emergence were also determined. Den entry for females began during the fourth week in September, with 90% denned by the fourth week of November. Earliest den entry for males occurred during the second week of October, with 90% denned by the second week of December. Mean week of den entry for known pregnant females was earlier than males. Earliest week of den entry for known pregnant females was earlier than other females and males. Earliest den emergence for males occurred during the first week of February, with 90% of males out of dens by the fourth week of April. Earliest den emergence for females occurred during the third week of March; by the first week of May, 90% of females had emerged. Male bears emerged from dens earlier than females. Denning period differed among classes and averaged 171 days for females that emerged from dens with cubs, 151 days for other females, and 131 days for males. Known pregnant females tended to den at higher elevations and, following emergence, remained at higher elevation until late May. Females with cubs remained relatively close (<3 km) to den sites until the last 2 weeks in May. Timing of denning events was similar to previous estimates for this and other grizzly bear populations in the southern Rocky Mountains.

Wyoming

Radiotracking large wilderness mammals: Integration of GPS and Argos technology

We tested 30 prototype global positioning system (GPS) radiocollars on brown bears ( Ursus arctos ) over a 3-year period on the Kenai Peninsula, Alaska. Collars were of 2 design types: GPS with an Argos (Argos Data collection and Location System) satellite uplink ( n= 19) and GPS unites where the data were stored on board ( n =10) for retrieval at a later date. All units also contained a conventional VHF (very high frequency) transmitter and weighed 1.7 kg. GPS-Argos united obtained 10-82% of expected GPS fixes, and fix rate declined significantly ( P <0.05) with time after deployment. Argos uplink success (proportion of successful transmissions of stored data) was linearly related to GPS fix rate ( r =0.91, P <0.001). Store-on-board units obtained significantly more successful fixes when compared with the GPS-Argos units ( t =-4.009, P <0.001). Fix success rate for deployed store-on-board collars ranged from 13-96%; because of the increased number of attempted fixes per day, these collars obtained fixes on 97% of days deployed. Accuracy of the GPS units was less than predicted by the NAVSTAR GPS technology using the course acquisition code. Reduced accuracy was likely a result of the proportion of 2-dimensional versus 3-dimensional fixes obtained, although we could not determine this statistic from recorded data. Increased overstory closure was the only variable measured that partially explained the reduced likelihood of a successful fix. Stem density, stem diameter, and overstory height measured within 3 m of the collar did not affect fix success. GPS fix success rates for collars attached to bears varied more and were lower than fix rates for stationary collars placed in various vegetation types, suggesting that the bear, terrain, and movement all influence both fix and uplink success rate. Application of this new technology to grizzly and brown bear research and comparisons to studies with moose ( Alces alces ) are discussed.

Alaska

Radio-tracking large wilderness mammals: integration of GPS and Argos technologies

We tested 30 prototype global positioning system (GPS) radiocollars on brown bears ( Ursus arctos ) over a 3-year period on the Kenai Peninsula, Alaska. Collars were of 2 design types: GPS units with an Argos (Argos Data collection and Location System) satellite uplink ( n = 19) and GPS units where the data were stored on board ( n = 10) for retrieval at a later date. All units also contained a conventional VHF (very high frequency) transmitter and weighed 1.7 kg. GPS-Argos units obtained 10-82% of expected GPS fixes, and fix rate declined significantly ( P < 0.05) with time after deployment. Argos uplink success (proportion of successful transmissions of stored data) was linearly related to GPS fix rate ( r = 0.91, P < 0.001). Storeon-board units obtained significantly more successful fixes when compared with the GPS-Argos units ( t = -4.009, P < 0.001). Fix success rate for deployed store-on-board collars ranged from 13-96%; because of the increased number of attempted fixes per day, these collars obtained fixes on 97% of days deployed. Accuracy of the GPS units was less than predicted by the NAVSTAR GPS technology using the course acquisition code. Reduced accuracy was likely a result of the proportion of 2-dimensional versus 3-dimensional fixes obtained, although we could not determine this statistic from recorded data. Increased overstory closure was the only variable measured that partially explained the reduced likelihood of a successful fix. Stem density, stem diameter, and overstory height measured within 3 m of the collar did not affect fix success. GPS fix success rates for collars attached to bears varied more and were lower than fix rates for stationary collars placed in various vegetation types, suggesting that the bear, terrain, and movement all influence both fix and uplink success rate. Application of this new technology to grizzly and brown bear research and comparisons to studies with moose ( Alces alces ) are discussed.

Alaska

Effects of sample size and sampling frequency on studies of brown bear home ranges and habitat use

We equipped 9 brown bears ( Ursus arctos ) on the Kenai Peninsula, Alaska, with collars containing both conventional very-high-frequency (VHF) transmitters and global positioning system (GPS) receivers programmed to determine an animal's position at 5.75-hr intervals. We calculated minimum convex polygon (MCP) and fixed and adaptive kernel home ranges for randomly-selected subsets of the GPS data to examine the effects of sample size on accuracy and precision of home range estimates. We also compared results obtained by weekly aerial radiotracking versus more frequent GPS locations to test for biases in conventional radiotracking data. Home ranges based on the MCP were 20-606 km 2 ( x = 201) for aerial radiotracking data ( n = 12-16 locations/bear) and 116-1,505 km 2 ( x = 522) for the complete GPS data sets ( n = 245-466 locations/bear). Fixed kernel home ranges were 34-955 km 2 ( x = 224) for radiotracking data and 16-130 km 2 ( x = 60) for the GPS data. Differences between means for radiotracking and GPS data were due primarily to the larger samples provided by the GPS data. Means did not differ between radiotracking data and equivalent-sized subsets of GPS data (P > 0.10). For the MCP, home range area increased and variability decreased asymptotically with number of locations. For the kernel models, both area and variability decreased with increasing sample size. Simulations suggested that the MCP and kernel models required >60 and >80 locations, respectively, for estimates to be both accurate (change in area <1%/additional location) and precise (CV < 50%). Although the radiotracking data appeared unbiased, except for the relationship between area and sample size, these data failed to indicate some areas that likely were important to bears. Our results suggest that the usefulness of conventional radiotracking data may be limited by potential biases and variability due to small samples. Investigators that use home range estimates in statistical tests should consider the effects of variability of those estimates. Use of GPS-equipped collars can facilitate obtaining larger samples of unbiased data and improve accuracy and precision of home range estimates.

Ursus