Carnivore space use behaviors reveal variation in responses to human land modification.

BACKGROUND: Spatial behavior, including home-ranging behaviors, habitat selection, and movement, can be extremely informative in estimating how animals respond to landscape heterogeneity. Responses in these spatial behaviors to features such as human land modification and resources can highlight a species' spatial strategy to maximize fitness and minimize mortality. These strategies can vary on spatial, temporal, and individual scales, and the combination of behaviors on these scales can lead to very different strategies among species. METHODS: Harnessing the variation present at these scales, we characterized how species may respond to stimuli in their environments ranging from broad- to fine-scale spatial responses to human modification in their environment. Using 15 bobcat-years and 31 coyote-years of GPS data from individuals inhabiting a landscape encompassing a range of human land modification, we evaluated the complexity of both species' responses to human modification on the landscape through their home range size, habitat selection, and functional response behaviors, accounting for annual, seasonal, and diel variation. RESULTS: Bobcats and coyotes used different strategies in response to human modification in their home ranges, with bobcats broadly expanding their home range with increases in human modification and displaying temporal consistency in functional response in habitat selection across both season and time of day. Meanwhile, coyotes did not expand their home ranges with increased human modification, but instead demonstrated fine-scale responses to human modification with habitat selection strategies that sometimes varied by time of day and season, paired with functional responses in selection behaviors. CONCLUSIONS: These differences in response to habitat, resources, and human modification between the two species highlighted the variation in spatial behaviors animals can use to exist in anthropogenic environments. Categorizing animal spatial behavior based on these spatiotemporal responses and individual variation can help in predicting how a species will respond to future change based on their current spatial behavior.

Estimating encounter-habitat relationships with scale-integrated resource selection functions.

Encounters between animals occur when animals are close in space and time. Encounters are important in many ecological processes including sociality, predation and disease transmission. Despite this, there is little theory regarding the spatial distribution of encounters and no formal framework to relate environmental characteristics to encounters. The probability of encounter could be estimated with resource selection functions (RSFs) by comparing locations where encounters occurred to available locations where they may have occurred, but this estimate is complicated by the hierarchical nature of habitat selection. We developed a method to relate resources to the relative probability of encounter based on a scale-integrated habitat selection framework. This framework integrates habitat selection at multiple scales to obtain an appropriate estimate of availability for encounters. Using this approach, we related encounter probabilities to landscape resources. The RSFs describe habitat associations at four scales, home ranges within the study area, areas of overlap within home ranges, locations within areas of overlap, and encounters compared to other locations, which can be combined into a single scale-integrated RSF. We apply this method to intraspecific encounter data from two species: white-tailed deer (Odocoileus virginianus) and elk (Cervus elaphus) and interspecific encounter data from a two-species system of caribou (Rangifer tarandus) and coyote (Canis latrans). Our method produced scale-integrated RSFs that represented the relative probability of encounter. The predicted spatial distribution of encounters obtained based on this scale-integrated approach produced distributions that more accurately predicted novel encounters than a naïve approach or any individual scale alone. Our results highlight the importance of accounting for the conditional nature of habitat selection in estimating the habitat associations of animal encounters as opposed to 'naïve' comparisons of encounter locations with general availability. This method has direct relevance for testing hypotheses about the relationship between habitat and social or predator-prey behaviour and generating spatial predictions of encounters. Such spatial predictions may be vital for understanding the distribution of encounters driving disease transmission, predation rates and other population and community-level processes.

Habitat quality influences trade-offs in animal movement along the exploration-exploitation continuum.

To successfully establish itself in a novel environment, an animal must make an inherent trade-off between knowledge accumulation and exploitation of knowledge gained (i.e., the exploration-exploitation dilemma). To evaluate how habitat quality affects the spatio-temporal scale of switching between exploration and exploitation during home range establishment, we conducted experimental trials comparing resource selection and space-use of translocated animals to those of reference individuals using reciprocal translocations between habitat types of differing quality. We selected wild pigs (Sus scrofa) as a model species to investigate hypotheses related to the movement behavior of translocated individuals because they are globally distributed large mammals that are often translocated within their introduced range to facilitate recreational hunting. Individuals translocated to higher quality habitat (i.e. higher proportions of bottomland hardwood habitats) exhibited smaller exploratory movements and began exploiting resources more quickly than those introduced to lower quality areas, although those in lower-quality areas demonstrated an increased rate of selection for preferred habitat as they gained knowledge of the landscape. Our data demonstrate that habitat quality mediates the spatial and temporal scale at which animals respond behaviorally to novel environments, and how these processes may determine the success of population establishment.

Seasonal variation in space use and territoriality in a large mammal (Sus scrofa).

An individual's spatial behavior is shaped by social and environmental factors and provides critical information about population processes to inform conservation and management actions. Heterogeneity in spatial overlap among conspecifics can be evaluated using estimates of home ranges and core areas and used to understand factors influencing space use and territoriality. To understand and test predictions about spatial behavior in an invasive large mammal, the wild pig (Sus scrofa), we examined variation in space use between sexes and seasons. We predicted that if animals were territorial that there would be a reduction in space-use overlap when comparing overlap of home ranges (HR-HR), to home ranges and core areas (HR-CA), and in-turn between core areas (CA-CA). Home ranges and core areas were estimated for 54 wild pigs at Buck Island Ranch, FL from GPS telemetry data. Overlap indices were calculated to estimate the strength (space-use overlap) and number of potential interactions within three wet seasons (June-October) and two dry seasons (December-April). Among sexes, home range size did not vary seasonally, and males exhibited larger home ranges compared to females (M = 10.36 ± 0.79 km2 (± SE), F = 3.21 ± 0.16 km2). Strength of overlap varied by season with wild pig home ranges overlapping more during the dry season. Males interacted with a greater number of individuals of both sexes, compared to females, and exhibited greater strength of overlap during the dry season. Consistent with our predictions, wild pigs appeared to exhibit territorial behavior, where strength of overlap decreased when comparing HR-HR to HR-CA and HR-CA to CA-CA. Our framework can be used to understand patterns of space use and territoriality in populations, which has important implications in understanding intraspecific interactions and population processes, such as how pathogens and parasites might spread within and among populations.

Food resources affect territoriality of invasive wild pig sounders with implications for control.

Interest in control methods for invasive wild pigs (Sus scrofa) has increased due to their range expansion, population growth, and an improved understanding of their destructive ecological and economic effects. Recent technological advances in traps for control of pig populations facilitate capture of entire social groups (sounders), but the efficacy of "whole-sounder" trapping strategies is heavily dependent on the degree of territoriality among sounders, a topic little research has explored. We assessed territoriality in wild pig sounders on the Savannah River Site, South Carolina, USA, and examined whether availability of food resources provided by a municipal-waste landfill affected among-sounder territoriality. We estimated utilization distribution overlap and dynamic interactions among 18 neighboring sounders around a landfill. We found that although neighboring sounders overlapped in space, intensity of use in shared areas was uniformly low, indicating territorial behavior. Neighbors tended to share slightly more space when closer to the landfill waste cells, indicating availability of a super-abundant resource somewhat weakens the degree of territoriality among sounders. Nevertheless, we conclude that sounders behaved in a generally territorial manner, and we discuss implications for whole-sounder trapping programs, particularly near concentrated resources such as landfills and crop fields.

Multi-level movement response of invasive wild pigs (Sus scrofa) to removal.

BACKGROUND: Lethal removal of invasive species, such as wild pigs (Sus scrofa), is often the most efficient approach for reducing their negative impacts. Wild pigs are one of the most widespread and destructive invasive mammals in the USA. Lethal management techniques are a key approach for wild pigs and can alter wild pig spatial behavior, but it is unclear how wild pigs respond to the most common removal technique, trapping. We investigated the spatial behavior of wild pigs following intensive removal of conspecifics via trapping at three sites within the Savannah River Site, SC, USA. We evaluated changes in wild pig densities, estimated temporal shifts in home-range properties, and evaluated fine-scale movement responses of wild pigs to removal. RESULTS: We observed a significant reduction in the density of wild pigs in one site following removal via trapping while a qualitative reduction was observed in another site. We found little evidence of shifts in pig home-ranging behavior following removal. However, we did observe a nuanced response in movement behavior of wild pigs to the removal at the scale of the GPS locations (4 h), including increased movement speed and reduced selection for vegetation rich areas. CONCLUSION: Our work provides a better understanding of the impact of removal via trapping on wild pig movement and its implications for management. The lack of shift in home-range characteristics observed illustrates how targeted trapping could be used to provide temporary relief for species sensitive to wild pig consumption such as ground nesting birds or agricultural crops.

How spatio-temporal habitat connectivity affects amphibian genetic structure.

Heterogeneous landscapes and fluctuating environmental conditions can affect species dispersal, population genetics, and genetic structure, yet understanding how biotic and abiotic factors affect population dynamics in a fluctuating environment is critical for species management. We evaluated how spatio-temporal habitat connectivity influences dispersal and genetic structure in a population of boreal chorus frogs (Pseudacris maculata) using a landscape genetics approach. We developed gravity models to assess the contribution of various factors to the observed genetic distance as a measure of functional connectivity. We selected (a) wetland (within-site) and (b) landscape matrix (between-site) characteristics; and (c) wetland connectivity metrics using a unique methodology. Specifically, we developed three networks that quantify wetland connectivity based on: (i) P. maculata dispersal ability, (ii) temporal variation in wetland quality, and (iii) contribution of wetland stepping-stones to frog dispersal. We examined 18 wetlands in Colorado, and quantified 12 microsatellite loci from 322 individual frogs. We found that genetic connectivity was related to topographic complexity, within- and between-wetland differences in moisture, and wetland functional connectivity as contributed by stepping-stone wetlands. Our results highlight the role that dynamic environmental factors have on dispersal-limited species and illustrate how complex asynchronous interactions contribute to the structure of spatially-explicit metapopulations.