Jon S. Horne

Analyzing animal movements using Brownian bridges.

By studying animal movements, researchers can gain insight into many of the ecological characteristics and processes important for understanding population-level dynamics. We developed a Brownian bridge movement model (BBMM) for estimating the expected movement path of an animal, using discrete location data obtained at relatively short time intervals. The BBMM is based on the properties of a conditional random walk between successive pairs of locations, dependent on the time between locations, the distance between locations, and the Brownian motion variance that is related to the animals mobility. We describe two critical developments that enable widespread use of the BBMM, including a derivation of the model when location data are measured with error and a maximum likelihood approach for estimating the Brownian motion variance. After the BBMM is fitted to location data, an estimate of the animals probability of occurrence can be generated for an area during the time of observation. To illustrate potential applications, we provide three examples: estimating animal home ranges, estimating animal migration routes, and evaluating the influence of fine-scale resource selection on animal movement patterns.

Likelihood Cross-Validation Versus Least Squares Cross-Validation for Choosing the Smoothing Parameter in Kernel Home-Range Analysis

Abstract Fixed kernel density analysis with least squares cross-validation (LSCVh) choice of the smoothing parameter is currently recommended for home-range estimation. However, LSCVh has several drawbacks, including high variability, a tendency to undersmooth data, and multiple local minima in the LSCVh function. An alternative to LSCVh is likelihood cross-validation (CVh). We used computer simulations to compare estimated home ranges using fixed kernel density with CVh and LSCVh to true underlying distributions. Likelihood cross-validation generally performed better than LSCVh, producing estimates with better fit and less variability, and it was especially beneficial at sample sizes <˜50. Because CVh is based on minimizing the Kullback-Leibler distance and LSCVh the integrated squared error, for each of these measures of discrepancy, we discussed their foundation and general use, statistical properties as they relate to home-range analysis, and the biological or practical interpretation of these statistical properties. We found 2 important problems related to computation of kernel home-range estimates, including multiple minima in the LSCVh and CVh functions and discrepancies among estimates from current home-range software. Choosing an appropriate smoothing parameter is critical when using kernel methods to estimate animal home ranges, and our study provides useful guidelines when making this decision.

Identifying and prioritizing ungulate migration routes for landscape‐level conservation

As habitat loss and fragmentation increase across ungulate ranges, identifying and prioritizing migration routes for conservation has taken on new urgency. Here we present a general framework using the Brownian bridge movement model (BBMM) that: (1) provides a probabilistic estimate of the migration routes of a sampled population, (2) distinguishes between route segments that function as stopover sites vs. those used primarily as movement corridors, and (3) prioritizes routes for conservation based upon the proportion of the sampled population that uses them. We applied this approach to a migratory mule deer (Odocoileus hemionus) population in a pristine area of southwest Wyoming, USA, where 2000 gas wells and 1609 km of pipelines and roads have been proposed for development. Our analysis clearly delineated where migration routes occurred relative to proposed development and provided guidance for on-the-ground conservation efforts. Mule deer migration routes were characterized by a series of stopover sites where deer spent most of their time, connected by movement corridors through which deer moved quickly. Our findings suggest management strategies that differentiate between stopover sites and movement corridors may be warranted. Because some migration routes were used by more mule deer than others, proportional level of use may provide a reasonable metric by which routes can be prioritized for conservation. The methods we outline should be applicable to a wide range of species that inhabit regions where migration routes are threatened or poorly understood.

SELECTING THE BEST HOME RANGE MODEL: AN INFORMATION-THEORETIC APPROACH

Choosing an appropriate home range model is important for describing space use by animals and understanding the ecological processes affecting animal movement. Traditional approaches for choosing among home range models have not resulted in general, consistent, and unambiguous criteria that can be applied to individual data sets. We present a new application of information-theoretic model selection that overcomes many of the limitations of traditional approaches, as follows. (1) It alleviates the need to know the true home range to assess home range models, thus allowing performance to be evaluated with data on individual animals. (2) The best model can be chosen from a set of candidate models with the proper balance between fit and complexity. (3) If candidate home range models are based on underlying ecological processes, researchers can use the selected model not only to describe the home range, but also to infer the importance of various ecological processes affecting animal movements within the home range.

Correcting Home-Range Models for Observation Bias

Abstract Home-range models implicitly assume equal observation rates across the study area. Because this assumption is frequently violated, we describe methods for correcting home-range models for observation bias. We suggest corrections for 3 general types of home-range models including those for which parameters are estimated using least-squares theory, models utilizing maximum likelihood for parameter estimation, and models based on kernel smoothing techniques. When applied to mule deer (Odocoileus hemionus) location data, we found that uncorrected estimates of the utilization distribution were biased low by as much as 18.4% and biased high by 19.2% when compared to corrected estimates. Because the magnitude of bias is related to several factors, future research should determine the relative influence of each of these factors on home-range bias.

Identifying polar bear resource selection patterns to inform offshore development in a dynamic and changing Arctic

Although sea ice loss is the primary threat to polar bears (Ursus maritimus), little can be done to mitigate its effects without global efforts to reduce greenhouse gas emissions. Other factors, however, could exacerbate the impacts of sea ice loss on polar bears, such as exposure to increased industrial activity. The Arctic Ocean has enormous oil and gas potential, and its development is expected to increase in the coming decades. Estimates of polar bear resource selection will inform managers how bears use areas slated for oil development and to help guide conservation planning. We estimated temporally-varying resource selection patterns for non-denning adult female polar bears in the Chukchi Sea population (2008–2012) at two scales (i.e., home range and weekly steps) to identify factors predictive of polar bear use throughout the year, before any offshore development. From the best models at each scale, we estimated scale-integrated resource selection functions to predict polar bear space use across th...

Habitat Partitioning by Sympatric Ocelots and Bobcats: Implications for Recovery of Ocelots in Southern Texas

Abstract Populations of ocelots (Leopardus pardalis) have declined during the past century due mainly to loss of habitat resulting in the ocelot being listed as endangered by the United States Fish and Wildlife Service. In southern Texas, the northern distribution of the ocelot overlaps the southern distribution of the bobcat (Lynx rufus). Because bobcats could adversely affect populations of ocelots through interspecific competition, we examined habitat selection of sympatric ocelots and bobcats to determine if habitat partitioning could be functioning to reduce interspecific interactions. Using radiotelemetry, we analyzed macro-scale (vegetative communities) and micro-scale (structural components) selection of habitats by sympatric ocelots and bobcats on Laguna Atascosa National Wildlife Refuge, Cameron County, Texas. We looked for differences in placement of home ranges within the general study area, selection of cover within home ranges, and use of structural components of vegetation within types of cover. There was substantial evidence for habitat partitioning with ocelots selecting areas with >75% canopy cover, while bobcats selected areas with <75% canopy cover. Thus, coexistence between these two species might be facilitated by resource partitioning of habitat.

Modeling Caribou Movements: Seasonal Ranges and Migration Routes of the Central Arctic Herd.

Migration is an important component of the life history of many animals, but persistence of large-scale terrestrial migrations is being challenged by environmental changes that fragment habitats and create obstacles to animal movements. In northern Alaska, the Central Arctic herd (CAH) of barren-ground caribou (Rangifer tarandus granti) is known to migrate over large distances, but the herd’s seasonal distributions and migratory movements are not well documented. From 2003–2007, we used GPS radio-collars to determine seasonal ranges and migration routes of 54 female caribou from the CAH. We calculated Brownian bridges to model fall and spring migrations for each year and used the mean of these over all 4 years to identify areas that were used repeatedly. Annual estimates of sizes of seasonal ranges determined by 90% fixed kernel utilization distributions were similar between summer and winter (X̅ = 27,929 SE = 1,064 and X̅ = 26,585 SE = 4912 km2, respectively). Overlap between consecutive summer and winter ranges varied from 3.3–18.3%. Percent overlap between summer ranges used during consecutive years (X̅ = 62.4% SE = 3.7%) was higher than for winter ranges (X̅ = 42.8% SE = 5.9%). Caribou used multiple migration routes each year, but some areas were used by caribou during all years, suggesting that these areas should be managed to allow for continued utilization by caribou. Restoring migration routes after they have been disturbed or fragmented is challenging. However, prior knowledge of movements and threats may facilitate maintenance of migratory paths and seasonal ranges necessary for long-term persistence of migratory species.

Quantifying the importance of patch‐specific changes in habitat to metapopulation viability of an endangered songbird

A growing number of programs seek to facilitate species conservation using incentive-based mechanisms. Recently, a market-based incentive program for the federally endangered Golden-cheeked Warbler (Dendroica chrysoparia) was implemented on a trial basis at Fort Hood, an Army training post in Texas, USA. Under this program, recovery credits accumulated by Fort Hood through contracts with private landowners are used to offset unintentional loss of breeding habitat of Golden-cheeked Warblers within the installation. Critical to successful implementation of such programs is the ability to value, in terms of changes to overall species viability, both habitat loss and habitat restoration or protection. In this study, we sought to answer two fundamental questions: Given the same amount of change in breeding habitat, does the change in some patches have a greater effect on metapopulation persistence than others? And if so, can characteristics of a patch (e.g., size or spatial location) be used to predict how the metapopulation will respond to these changes? To answer these questions, we describe an approach for using sensitivity analysis of a metapopulation projection model to predict how changes to specific habitat patches would affect species viability. We used a stochastic, discrete-time projection model based on stage-specific estimates of survival and fecundity, as well as various assumptions about dispersal among populations. To assess a particular patchs leverage, we quantified how much metapopulation viability was expected to change in response to changing the size of that patch. We then related original patch size and distance from the largest patch to each patchs leverage to determine if general patch characteristics could be used to develop guidelines for valuing changes to patches within a metapopulation. We found that both the characteristic that best predicted patch leverage and the magnitude of the relationship changed under different model scenarios. Thus, we were unable to find a consistent set of relationships, and therefore we emphasize the dangers in relying on general guidelines to assess patch value. Instead, we provide an approach that can be used to quantitatively evaluate patch value and identify critical needs for future research.