Application of radiotelemetry to wildlife conservation in mountainous Asian landscapes
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Book chapter↗
Geology topics
M.S. Mitchell
Publications and source records attributed to M.S. Mitchell.
Wolf population dynamics in the U.S. Northern Rocky Mountains are affected by recruitment and human-caused mortality
Reliable analyses can help wildlife managers make good decisions, which are particularly critical for controversial decisions such as wolf (Canis lupus) harvest. Creel and Rotella (2010) recently predicted substantial population declines in Montana wolf populations due to harvest, in contrast to predictions made by Montana Fish, Wildlife and Parks (MFWP). We replicated their analyses considering only those years in which field monitoring was consistent, and we considered the effect of annual variation in recruitment on wolf population growth. Rather than assuming constant rates, we used model selection methods to evaluate and incorporate models of factors driving recruitment and human-caused mortality rates in wolf populations in the Northern Rocky Mountains. Using data from 27 area-years of intensive wolf monitoring, we show that variation in both recruitment and human-caused mortality affect annual wolf population growth rates and that human-caused mortality rates have increased with the sizes of wolf populations. We document that recruitment rates have decreased over time, and we speculate that rates have decreased with increasing population sizes and/or that the ability of current field resources to document recruitment rates has recently become less successful as the number of wolves in the region has increased. Estimates of positive wolf population growth in Montana from our top models are consistent with field observations and estimates previously made by MFWP for 2008–2010, whereas the predictions for declining wolf populations of Creel and Rotella (2010) are not. Familiarity with limitations of raw data, obtained first-hand or through consultation with scientists who collected the data, helps generate more reliable inferences and conclusions in analyses of publicly available datasets. Additionally, development of efficient monitoring methods for wolves is a pressing need, so that analyses such as ours will be possible in future years when fewer resources will be available for monitoring.
Idaho, Montana, Wyoming↗
Estimation of successful breeding pairs for wolves in the Northern Rocky Mountains, USA
Under the Endangered Species Act, documenting recovery and federally mandated population levels of wolves (Canis lupus) in the Northern Rocky Mountains (NRM) requires monitoring wolf packs that successfully recruit young. United States Fish and Wildlife Service regulations define successful breeding pairs as packs estimated to contain an adult male and female, accompanied by ???2 pups on 31 December of a given year. Monitoring successful breeding pairs will become more difficult following proposed delisting of NRM wolves; alternatives to historically intensive methods, appropriate to the different ecological and regulatory context following delisting, are required. Because pack size is easier to monitor than pack composition, we estimated probability a pack would contain a successful breeding pair based on its size for wolf populations inhabiting 6 areas in the NRM. We also evaluated the extent to which differences in demography of wolves and levels of human-caused mortality among the areas influenced the probability of packs of different sizes would contain successful breeding pairs. Probability curves differed among analysis areas, depending primarily on levels of human-caused mortality, secondarily on annual population growth rate, and little on annual population density. Probabilities that packs contained successful breeding pairs were more uniformly distributed across pack sizes in areas with low levels of human mortality and stable populations. Large packs in areas with high levels of human-caused mortality and high annual growth rates had relatively high probabilities of containing breeding pairs whereas those for small packs were relatively low. Our approach can be used by managers to estimate number of successful breeding pairs in a population where number of packs and their sizes are known. Following delisting of NRM wolves, human-caused mortality is likely to increase, resulting in more small packs with low probabilities of containing breeding pairs. Differing contributions of packs to wolf population growth based on their size suggests monitoring successful breeding pairs will provide more accurate insights into population dynamics of wolves than will monitoring number of packs or individuals only.
Journal of Wildlife Management↗
Territoriality of feral pigs in a highly persecuted population on Fort Benning, Georgia
We examined home range behavior of female feral pigs (Sus scrofa) in a heavily hunted population on Fort Benning Military Reservation in west-central Georgia, USA. We used Global Positioning System location data from 24 individuals representing 18 sounders (i.e., F social groups) combined with markrecapture and camera-trap data to evaluate evidence of territorial behavior at the individual and sounder levels. Through a manipulative experiment, we examined evidence for an inverse relationship between population density and home range size that would be expected for territorial animals. Pigs from the same sounder had extensive home range overlap and did not have exclusive core areas. Sounders had nearly exclusive home ranges and had completely exclusive core areas, suggesting that female feral pigs on Fort Benning were territorial at the sounder level but not at the individual level. Lethal removal maintained stable densities of pigs in our treatment area, whereas density increased in our control area; territory size in the 2 areas was weakly and inversely related to density of pigs. Territorial behavior in feral pigs could influence population density by limiting access to reproductive space. Removal strategies that 1) match distribution of removal efforts to distribution of territories, 2) remove entire sounders instead of individuals, and 3) focus efforts where high-quality food resources strongly influence territorial behaviors may be best for long-term control of feral pigs.
Journal of Wildlife Management↗
Estimate of herpetofauna depredation by a population of wild pigs
Herpetofauna populations are decreasing worldwide, and the range of wild pigs (Sus scrofa) is expanding. Depredation of threatened reptile and amphibian populations by wild pigs could be substantial. By understanding depredation characteristics and rates, more resources can be directed toward controlling populations of wild pigs coincident with threatened or endangered herpetofauna populations. From April 2005 to March 2006 we used firearms to collect wild pigs (n = 68) and examined stomach content for reptiles and amphibians. We found 64 individual reptiles and amphibians, composed of 5 different species, that were consumed by wild pigs during an estimated 254 hours of foraging. Primarily arboreal species (e.g., Anolis carolinensis) became more vulnerable to depredation when temperatures were low and they sought thermal shelter. Other species (e.g., Scaphiopus holbrookii) that exhibit mass terrestrial migrations during the breeding season also faced increased vulnerability to depredation by wild pigs. Results suggest that wild pigs are opportunistic consumers that can exploit and potentially have a negative impact on species with particular life-history characteristics. ?? 2009 American Society of Mammalogists.
Journal of Mammalogy↗
Effect of experimental manipulation on survival and recruitment of feral pigs
Lethal removal is commonly used to reduce the density of invasive-species populations, presuming it reduces population growth rate; the actual effect of lethal removal on the vital rates contributing to population growth, however, is rarely tested. We implemented a manipulative experiment of feral pig (Sus scrofa) populations at Fort Benning, Georgia, USA, to assess the demographic effects of harvest intensity. Using markrecapture data, we estimated annual survival, recruitment, and population growth rates of populations in a moderately harvested area and a heavily harvested area for 200406. Population growth rates did not differ between the populations. The top-ranked model for survival included a harvest intensity effect; model-averaged survival was lower for the heavily harvested population than for the moderately harvested population. Increased immigration and reproduction likely compensated for the increased mortality in the heavily harvested population. We conclude that compensatory responses in feral pig recruitment can limit the success of lethal control efforts. ?? 2009 CSIRO.
Wildlife Research↗
Sources, sinks, and spatial ecology of cotton mice in longleaf pine stands undergoing restoration
The Fire and Fire Surrogate studya replicated, manipulative experimentsought the most economically and ecologically efficient way to restore the nation's fire-maintained ecosystems. As part of this study, we conducted a 3-year markrecapture study, comprising 105,000 trap-nights, to assess demographic responses of cotton mice (Peromyscus gossypinus) to Fire and Fire Surrogate treatments at the Gulf Coastal Plain site, where longleaf pine was the ecosystem to be restored. We compared competing models to evaluate restoration effects on variation in apparent survival and recruitment over time, space, and treatment, and incorporated measures of available source habitat for cotton mice with reverse-time modeling to infer immigration from outside the study area. The top-ranked survival model contained only variation over time, but the closely ranked 2nd and 3rd models included variation over space and treatment, respectively. The top 4 recruitment models all included effects for availability of source habitat and treatments. Burning appeared to degrade habitat quality for cotton mice, showing demographic characteristics of a sink, but treatments combining fire with thinning of trees or application of herbicide to the understory appeared to improve habitat quality, possibly creating sources. Bottomland hardwoods outside the study also acted as sources by providing immigrants to experimental units. Models suggested that population dynamics operated over multiple spatial scales. Treatments applied to 15-ha stands probably only caused local variation in vital rates within the larger population. ?? 2009 American Society of Mammalogists.
Journal of Mammalogy↗
Contributions of vital rates to growth of a protected population of American black bears
Analyses of large, long-lived animals suggest that adult survival generally has the potential to contribute more than reproduction to population growth rate (λ), but because survival varies little, high variability in reproduction can have a greater influence. This pattern has been documented for several species of large mammals, but few studies have evaluated such contributions of vital rates to λ for American black bears ( Ursus americanus ). We used variance-based perturbation analyses (life table response experiments, LTRE) and analytical sensitivity and elasticity analyses to examine the actual and potential contributions of variation of vital rates to variation in growth rate (λ) of a population of black bears inhabiting the Pisgah Bear Sanctuary in the southern Appalachian Mountains of North Carolina, using a 22-year dataset. We found that recruitment varied more than other vital rates; LTRE analyses conducted over several time intervals thus indicated that recruitment generally contributed at least as much as juvenile and adult survival to observed variation in λ, even though the latter 2 vital rates had the greater potential to affect λ. Our findings are consistent with predictions from studies on polar bears ( U. maritimus ) and grizzly bears ( U. arctos ), but contrast with the few existing studies on black bears in ways that suggest levels of protection from human-caused mortality might explain whether adult survival or recruitment contribute most to variation in λ for this species. We hypothesize that λ is most strongly influenced by recruitment in protected populations where adult survival is relatively high and constant, whereas adult survival will most influence λ for unprotected populations.
Ursus↗
Evaluating intercepts from demographic models to understand resource limitation and resource thresholds
Understanding resource limitation is critical to effective management and conservation of wild populations, however resource limitation is difficult to quantify partly because resource limitation is a dynamic process. Specifically, a resource that is limiting at one time may become non-limiting at another time, depending upon changes in its availability and changes in the availability of other resources. Methods for understanding resource limitation, therefore, must consider the dynamic effects of resources on demography. We present approaches for interpreting results of demographic modeling beyond analyzing model rankings, model weights, slope estimates, and model averaging. We demonstrate how interpretation of y-intercepts, odds ratios, and rates of change can yield insights into resource limitation as a dynamic process, assuming logistic regression is used to link estimates of resources with estimates of demography. In addition, we show how x-intercepts can be evaluated with respect to odds ratios to understand resource thresholds. ?? 2007 Elsevier B.V. All rights reserved.
Ecological Modelling↗
Estimated home ranges can misrepresent habitat relationships on patchy landscapes
Home ranges of animals are generally structured by the selective use of resource-bearing patches that comprise habitat. Based on this concept, home ranges of animals estimated from location data are commonly used to infer habitat relationships. Because home ranges estimated from animal locations are largely continuous in space, the resource-bearing patches selected by an animal from a fragmented distribution of patches would be difficult to discern; unselected patches included in the home range estimate would bias an understanding of important habitat relationships. To evaluate potential for this bias, we generated simulated home ranges based on optimal selection of resource-bearing patches across a series of simulated resource distributions that varied in the spatial continuity of resources. For simulated home ranges where selected patches were spatially disjunct, we included interstitial, unselected cells most likely to be traveled by an animal moving among selected patches. We compared characteristics of the simulated home ranges with and without interstitial patches to evaluate how insights derived from field estimates can differ from actual characteristics of home ranges, depending on patchiness of landscapes. Our results showed that contiguous home range estimates could lead to misleading insights on the quality, size, resource content, and efficiency of home ranges, proportional to the spatial discontinuity of resource-bearing patches. We conclude the potential bias of including unselected, largely irrelevant patches in the field estimates of home ranges of animals can be high, particularly for home range estimators that assume uniform use of space within home range boundaries. Thus, inferences about the habitat relationships that ultimately define an animal's home range can be misleading where animals occupy landscapes with patchily distributed resources.
Ecological Modelling↗
Projected long-term response of Southeastern birds to forest management
Numerous studies have explored the influence of forest management on avian communities empirically, but uncertainty about causal relationships between landscape patterns and temporal dynamics of bird communities calls into question how observed historical patterns can be projected into the future, particularly to assess consequences of differing management alternatives. We used the Habplan harvest scheduler to project forest conditions under several management scenarios mapped at 5-year time steps over a 40-year time span. We used empirical models of overall avian richness, richness of selected guilds, and probability of presence for selected species to predict avian community characteristics for each of the mapped landscapes generated for each 5-year time step for each management scenario. We then used time series analyses to quantify relationships between changes in avian community characteristics and management-induced changes to forest landscapes over time. Our models of avian community and species characteristics indicated habitat associations at multiple spatial scales, although landscape-level measures of habitat were generally more important than stand-level measures. Our projections showed overall avian richness, richness of Neotropical migrants, and the presence of Blue-gray Gnatcatchers and Eastern Wood-pewees varied little among management scenarios, corresponding closely to broad, overall landscape changes over time. By contrast, richness of canopy nesters, richness of cavity nesters, richness of scrub-successional associates, and the presence of Common Yellowthroats showed high temporal variability among management scenarios, likely corresponding to short-term, fine-scale changes in the landscape. Predicted temporal variability of both interior-forest and early successional birds was low in the unharvested landscape relative to that in the harvested landscape. Our results also suggested that early successional species can be sensitive to both availability and connectivity of habitat on the landscape. To increase or maintain the avian diversity, our projections indicate that forest managers need to consider landscape-scale configuration of stands, maintaining a spatially heterogeneous distribution of age classes. Our findings suggest which measures of richness or species presence may be appropriate indicators for monitoring effects of forest management on avian communities, depending on management objectives.
Forest Ecology and Management↗
Change-in-ratio density estimator for feral pigs is less biased than closed mark-recapture estimates
Closed-population capture-mark-recapture (CMR) methods can produce biased density estimates for species with low or heterogeneous detection probabilities. In an attempt to address such biases, we developed a density-estimation method based on the change in ratio (CIR) of survival between two populations where survival, calculated using an open-population CMR model, is known to differ. We used our method to estimate density for a feral pig (Sus scrofa) population on Fort Benning, Georgia, USA. To assess its validity, we compared it to an estimate of the minimum density of pigs known to be alive and two estimates based on closed-population CMR models. Comparison of the density estimates revealed that the CIR estimator produced a density estimate with low precision that was reasonable with respect to minimum known density. By contrast, density point estimates using the closed-population CMR models were less than the minimum known density, consistent with biases created by low and heterogeneous capture probabilities for species like feral pigs that may occur in low density or are difficult to capture. Our CIR density estimator may be useful for tracking broad-scale, long-term changes in species, such as large cats, for which closed CMR models are unlikely to work. ?? CSIRO 2008.
Wildlife Research↗
Effects of roads on habitat quality for bears in the southern Appalachians: A long-term study
We tested the hypothesis that gravel roads, not paved roads, had the largest negative effect on habitat quality for a population of American black bears (Ursus americanus) that lived in a protected area, where vehicle collision was a relatively minimal source of mortality. We also evaluated whether road use by bears differed by sex or age and whether annual variation in hard mast productivity affected the way bears used areas near roads. In addition, we tested previous findings regarding the spatial extent to which roads affected bear behavior negatively. Using summer and fall home ranges for 118 black bears living in the Pisgah Bear Sanctuary in western North Carolina during 1981-2001, we estimated both home-range-scale (2nd-order) and within-home-range-scale (3rd-order) selection for areas within 250, 500, 800, and 1,600 m of paved and gravel roads. All bears avoided areas near gravel roads more than they avoided areas near paved roads during summer and fall for 2nd-order selection and during summer for 3rd-order selection. During fall, only adult females avoided areas near gravel roads more than they avoided areas near paved roads for 3rd-order selection. We found a positive relationship between use of roads by adults and annual variability in hard mast productivity. Overall, bears avoided areas within 800 m of gravel roads. Future research should determine whether avoidance of gravel roads by bears affects bear survival. ?? 2007 American Society of Mammalogists.
Journal of Mammalogy↗
Selection of den sites by black bears in the southern Appalachians
We evaluated selection of den sites by American black bears (Ursus americanus) in the Pisgah Bear Sanctuary, western North Carolina, by comparing characteristics of dens at 53 den sites with availability of habitat characteristics in annual home ranges of bears and in the study area. We also tested whether den-site selection differed by sex, age, and reproductive status of bears. In addition, we evaluated whether the den component of an existing habitat model for black bears predicted where bears would select den sites. We found bears selected den sites far from gravel roads, on steep slopes, and at high elevations relative to what was available in both annual home ranges and in the study area. Den-site selection did not differ by sex or age, but it differed by reproductive status. Adult females with cubs preferred to den in areas that were relatively far from gravel roads, but adult females without cubs did not. The habitat model overestimated the value of areas near gravel roads, underestimated the value of moderately steep areas, and did not include elevation as a predictor variable. Our results highlight the importance of evaluating den selection in terms of both use and availability of den characteristics. ?? 2007 American Society of Mammalogists.
Journal of Mammalogy↗
Linking resources with demography to understand resource limitation for bears
1. Identifying the resources that limit growth of animal populations is essential for effective conservation; however, resource limitation is difficult to quantify. Recent advances in geographical information systems (GIS) and resource modelling can be combined with demographic modelling to yield insights into resource limitation. 2. Using long-term data on a population of black bears Ursus americanus, we evaluated competing hypotheses about whether availability of hard mast (acorns and nuts) or soft mast (fleshy fruits) limited bears in the southern Appalachians, USA, during 1981-2002. The effects of clearcutting on habitat quality were also evaluated. Annual survival, recruitment and population growth rate were estimated using capture-recapture data from 101 females. The availability of hard mast, soft mast and clearcuts was estimated with a GIS, as each changed through time as a result of harvest and succession, and then availabilities were incorporated as covariates for each demographic parameter. 3. The model with the additive availability of hard mast and soft mast across the landscape predicted survival and population growth rate. Availability of young clearcuts predicted recruitment, but not population growth or survival. 4. Availability of hard mast stands across the landscape and availability of soft mast across the landscape were more important than hard mast production and availability of soft mast in young clearcuts, respectively. 5. Synthesis and applications. Our results indicate that older stands, which support high levels of hard mast and moderate levels of soft mast, should be maintained to sustain population growth of bears in the southern Appalachians. Simultaneously, the acreage of intermediate aged stands (10-25 years), which support very low levels of both hard mast and soft mast, should be minimized. The approach used in this study has broad application for wildlife management and conservation. State and federal wildlife agencies often possess long-term data on both resource availability and capture-recapture for wild populations. Combined, these two data types can be used to estimate survival, recruitment, population growth, elasticities of vital rates and the effects of resource availability on demographic parameters. Hence data that are traditionally used to understand population trends can be used to evaluate how and why demography changes over time. ?? 2007 The Authors.
Journal of Applied Ecology↗
Optimal use of resources structures home ranges and spatial distribution of black bears
Research has shown that territories of animals are economical. Home ranges should be similarly efficient with respect to spatially distributed resources and this should structure their distribution on a landscape, although neither has been demonstrated empirically. To test these hypotheses, we used home range models that optimize resource use according to resource-maximizing and area-minimizing strategies to evaluate the home ranges of female black bears, Ursus americanus, living in the southern Appalachian Mountains. We tested general predictions of our models using 104 home ranges of adult female bears studied in the Pisgah Bear Sanctuary, North Carolina, U.S.A., from 1981 to 2001. We also used our models to estimate home ranges for each real home range under a variety of strategies and constraints and compared similarity of simulated to real home ranges. We found that home ranges of female bears were efficient with respect to the spatial distribution of resources and were best explained by an area-minimizing strategy with moderate resource thresholds and low levels of resource depression. Although resource depression probably influenced the spatial distribution of home ranges on the landscape, levels of resource depression were too low to quantify accurately. Home ranges of lactating females had higher resource thresholds and were more susceptible to resource depression than those of breeding females. We conclude that home ranges of animals, like territories, are economical with respect to resources, and that resource depression may be the mechanism behind ideal free or ideal preemptive distributions on complex, heterogeneous landscapes. ?? 2007 The Association for the Study of Animal Behaviour.
Animal Behaviour↗
Relationships between avian richness and landscape structure at multiple scales using multiple landscapes
Little is known about factors that structure biodiversity on landscape scales, yet current land management protocols, such as forest certification programs, place an increasing emphasis on managing for sustainable biodiversity at landscape scales. We used a replicated landscape study to evaluate relationships between forest structure and avian diversity at both stand and landscape-levels. We used data on bird communities collected under comparable sampling protocols on four managed forests located across the Southeastern US to develop logistic regression models describing relationships between habitat factors and the distribution of overall richness and richness of selected guilds. Landscape models generated for eight of nine guilds showed a strong relationship between richness and both availability and configuration of landscape features. Diversity of topographic features and heterogeneity of forest structure were primary determinants of avian species richness. Forest heterogeneity, in both age and forest type, were strongly and positively associated with overall avian richness and richness for most guilds. Road density was associated positively but weakly with avian richness. Landscape variables dominated all models generated, but no consistent patterns in metrics or scale were evident. Model fit was strong for neotropical migrants and relatively weak for short-distance migrants and resident species. Our models provide a tool that will allow managers to evaluate and demonstrate quantitatively how management practices affect avian diversity on landscapes.
Forest Ecology and Management↗
A method for landscape analysis of forestry guidelines using bird habitat models and the Habplan harvest scheduler
Wildlife-habitat relationship models have sometimes been linked with forest simulators to aid in evaluating outcomes of forest management alternatives. However, linking wildlife-habitat models with harvest scheduling software would provide a more direct method for assessing economic and ecological implications of alternative harvest schedules in commercial forest operations. We demonstrate an approach for frontier analyses of wildlife benefits using the Habplan harvest scheduler and spatially explicit wildlife response models in the context of operational forest planning. We used the Habplan harvest scheduler to plan commercial forest management over a 40-year horizon at a landscape scale under five scenarios: unmanaged, an unlimited block-size option both with and without riparian buffers, three cases with different block-size restrictions, and a set-asides scenario in which older stands were withheld from cutting. The potential benefit to wildlife was projected based on spatial models of bird guild richness and species probability of detection. Harvested wood volume provided a measure of scenario costs, which provides an indication of management feasibility. Of nine species and guilds, none appeared to benefit from 50 m riparian buffers, response to an unmanaged scenario was mixed and expensive, and block-size restrictions (maximum harvest unit size) provided no apparent benefit and in some cases were possibly detrimental to bird richness. A set-aside regime, however, appeared to provide significant benefits to all species and groups, probably through increased landscape heterogeneity and increased availability of older forest. Our approach shows promise for evaluating costs and benefits of forest management guidelines in commercial forest enterprises and improves upon the state of the art by utilizing an optimizing harvest scheduler as in commercial forest management, multiple measures of biodiversity (models for multiple species and guilds), and spatially explicit wildlife response models. ?? 2006 Elsevier B.V. All rights reserved.
Forest Ecology and Management↗