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↗
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↗
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↗
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↗
Spatio-temporal availability of soft mast in clearcuts in the Southern Appalachians
Soft mast is an important resource for many wild populations in the Southern Appalachians, yet the way clear-cutting affects availability of soft mast though time is not fully understood. We tested a theoretical model of temporal availability of soft mast in clearcuts using empirical data on percent cover and berry production of Gaylussacia, Vaccinium, and Rubus spp. plants in 100 stands that were clearcut (0-122 years old) in the Southern Appalachian Mountains. We modeled the relationship between soft mast availability and stand age, evaluated the effects of topography and forest type on soft mast, developed statistical models for predicting the spatio-temporal distribution of soft mast, and tested the hypothesis that percent cover of berry plants and berry production provided similar information about soft mast availability. We found temporal dynamics explained berry production better than it predicted percent plant cover, whereas topographic variables influenced percent plant cover more than they influenced berry production. Berry production and percent plant cover were highest in ???2-9-year-old stands. Percent plant cover was lowest in 10-69-year-old stands and intermediate in 70+-year-old stands. Three of our spatio-temporal models performed well during model testing and they were not biased by the training data, indicating the inferences about spatio-temporal availability of soft mast extended beyond our sample data. The methods we used to estimate the distribution of soft mast may be useful for modeling distributions of other resources. ?? 2006 Elsevier B.V. All rights reserved.
Forest Ecology and Management↗
Long-term analysis of survival, fertility, and population growth rate of black bears in North Carolina
We estimated survival, fertility, and realized and asymptotic population growth rates from 1981 to 2002 for a protected population of black bears (Ursus americanus) in the southern Appalachian Mountains. We used Akaike's information criterion to assess the time interval for averaging observations that was best for estimating vital rates for our study, given our yearly sample sizes. The temporal symmetry approach allowed us to directly assess population growth and to address all losses and gains to the population by using only capture data, offering an alternative to the logistically intensive collection of reproductive data. Models that averaged survival and fertility across 5- and 7-year time intervals were best supported by our data. Studies of black bear populations with annual sample sizes similar to ours should be of at least 5 years in duration to estimate vital rates reliably, and at least 10 years in duration to evaluate changes in population growth rate (??). We also hypothesized that survival would not track changes in ?? because ?? is influenced by both survival and fertility. The 5-year model supported our hypothesis, but the 7-year model did not. Where long-term dynamics of large, relatively stable bear populations are of interest, monitoring survival is likely to be sufficient for evaluating trends in ??. For rapidly changing, small populations, however, failure to incorporate fertility into assessments of ?? could be misleading. ?? 2005 American Society of Mammalogists.
Journal of Mammalogy↗
Test of a habitat suitability index for black bears in the southern Appalachians
We present a habitat suitability index (HSI) model for black bears (Ursus americanus) living in the southern Appalachians that was developed a priori from the literature, then tested using location and home range data collected in the Pisgah Bear Sanctuary, North Carolina, over a 12-year period. The HSI was developed and initially tested using habitat and bear data collected over 2 years in the sanctuary. We increased number of habitat sampling sites, included data collected in areas affected by timber harvest, used more recent Geographic Information System (GIS) technology to create a more accurate depiction of the HSI for the sanctuary, evaluated effects of input variability on HSI values, and duplicated the original tests using more data. We found that the HSI predicted habitat selection by bears on population and individual levels and the distribution of collared bears were positively correlated with HSI values. We found a stronger relationship between habitat selection by bears and a second-generation HSI. We evaluated our model with criteria suggested by Roloff and Kernohan (1999) for evaluating HSI model reliability and concluded that our model was reliable and robust. The model's strength is that it was developed as an a priori hypothesis directly modeling the relationship between critical resources and fitness of bears and tested with independent data. We present the HSI spatially as a continuous fitness surface where potential contribution of habitat to the fitness of a bear is depicted at each point in space.
Wildlife Society Bulletin↗
Wildlife tradeoffs based on landscape models of habitat
It is becoming increasingly clear that the spatial structure of landscapes affects the habitat choices and abundance of wildlife. In contrast to wildlife management based on preservation of critical habitat features such as nest sites on a beach or mast trees, it has not been obvious how to incorporate spatial structure into management plans. We present techniques to accomplish this goal. We used multiscale logistic regression models developed previously for neotropical migrant bird species habitat use in South Carolina (USA) as a basis for these techniques. Based on these models we used a spatial optimization technique to generate optimal maps (probability of occurrence, P = 1.0) for each of seven species. To emulate management of a forest for maximum species diversity, we defined the objective function of the algorithm as the sum of probabilities over the seven species, resulting in a complex map that allowed all seven species to coexist. The map that allowed for coexistence is not obvious, must be computed algorithmically, and would be difficult to realize using rules of thumb for habitat management. To assess how management of a forest for a single species of interest might affect other species, we analyzed tradeoffs by gradually increasing the weighting on a single species in the objective function over a series of simulations. We found that as habitat was increasingly modified to favor that species, the probability of presence for two of the other species was driven to zero. This shows that whereas it is not possible to simultaneously maximize the likelihood of presence for multiple species with divergent habitat preferences, compromise solutions are possible at less than maximal likelihood in many cases. Our approach suggests that efficiency of habitat management for species diversity can by maximized for even small landscapes by incorporating spatial context. The methods we present are suitable for wildlife management, endangered species conservation, and nature reserve design.
Conference Paper↗