Home-range estimation within complex restricted environments: importance of method selection in detecting seasonal change

Context Despite the diversity of available home range estimators, no single method performs equally well in all circumstances. It is therefore important to understand how different estimators perform for data collected under diverse conditions. Kernel density estimation is a popular approach for home range estimation. While many studies have evaluated different kernel bandwidth selectors, few studies have compared different formulations of the bandwidth matrix using wildlife telemetry data. Additionally, few studies have compared the performance of kernel bandwidth selectors using VHF radio-telemetry data from small-bodied taxa. Aims In this study, we used eight different combinations of bandwidth selectors and matrices to evaluate their ability to meet several criteria that could be potentially used to select a home range estimator. Methods We used handheld VHF telemetry data from two species of snake displaying non-migratory and migratory movement patterns. We used subsampling to estimate each estimator’s sensitivity to sampling duration and fix rate and compared home range size, the number of disjunct volume contours and the proportion of telemetry fixes not included in those contours among estimators. Key Results We found marked differences among bandwidth selectors with regards to our criteria but comparatively little difference among bandwidth matrices for a given bandwidth selector. Least-squares cross-validation bandwidths exhibited near-universal convergence failure whereas likelihood cross-validation bandwidths showed high sensitivity to sampling duration and fix rate. The reference, plug-in and smoothed cross-validation bandwidths were more robust to variation in sampling intensity, with the former consistently producing the largest estimates of home range size. Conclusions Our study illustrates the performance of multiple kernel bandwidth estimators for estimating home ranges with datasets typical of many small-bodied taxa. The reference and plug-in bandwidths with an unconstrained bandwidth matrix generally had the best performance. However, our study concurs with earlier studies indicating that no single home range estimator performs equally well in all circumstances. Implications Although we did not find strong differences between bandwidth matrices, we encourage the use of unconstrained matrices because of their greater flexibility in smoothing data not parallel to the coordinate axes. We also encourage researchers to select an estimator suited to their study objectives and the life history of their study organism.

Radio-tracking squirrels: Performance of home range density and linkage estimators with small range and sample size

We compared the Fourier series, harmonic mean, minimum convex polygon, and 2 95% ellipse home-range estimators using computer-simulated data with a known home-range area. Data were generated to simulate home ranges with 1 or 2 centers of activity and with topographic barriers causing irregular shapes. Estimators performed with different precision and bias according to the type of data simulated and the number of observations taken. Overall, the harmonic mean was the least biased, but was also one of the least precise. Home-range estimators are only general measures of animal activity. J. WILDL. MANAGE. 54(2):310-315 Many statistical home-range estimators have been proposed. All have different underlying models (assumptions) providing different operating characteristics and therefore can produce dissimilar results for a specific set of data. Rarely have the statistical properties of these estimators been compared or have guidelines on the selection of a specific estimator for a particular set of data been proposed. Application of different estimators produces confusion in the interpretation of home-range estimates because some of the differences observed between studies are due to the estimators themselves, and not to the behavior of the animals being studied. We used computer simulated data to address the differences among estimators by comparing 3 nonparametric estimators: harmonic mean (Dixon and Chapman 1980), Fourier series (Anderson 1982), and minimum convex polygon (Mohr 1947); and 2 parametric estimators: 95% ellipse estimators of Jennrich and Turner (1969) and Koeppl et al. (1975). Two types of animal movements were simulated: (1) an animal with a center of activity (e.g., a nesting bird) and (2) an animal with a uniform distribution (no center of activity). Data from a normal distribution were used to simulate center-of-activity movements, and data from a uniform distribution were used for movements without a center of activity. In addition, home ranges with irregular This content downloaded from 207.46.13.114 on Thu, 26 May 2016 06:08:42 UTC All use subject to http://about.jstor.org/terms J. Wildl. Manage. 54(2):1990 HOME-RANGE ESTIMATORS * Boulanger and White 311 shapes, as would be caused by topographic barriers and/or habitat heterogeneity, were simulated. Advantages of using computer-simulated data are that a true home range is known, and thus bias and precision of each estimator can be evaluated. The statistical properties of an estimator cannot be determined with field telemetry data because true replicate observations cannot be constructed. Replicated, simulated data sets also allow powerful statistical comparisons of esti-

Home range estimators are a critical component for understanding animal spatial ecology. The choice of home range estimator in spatial ecology studies can significantly influence management and conservation actions, as different methods lead to vastly different interpretations of movement patterns, habitat selection, as well as home range requirements. Reptile studies in particular have struggled to reach a consensus on the appropriate home range estimators to use, and species with cryptic behavior make home range assessment difficult. We applied dynamic Brownian Bridge Movement Models (dBBMMs) to radio-telemetry data from Ophiophagus hannah, a wide-ranging snake species. We used two focal individuals at different life stages (one juvenile male and one adult male) and sought to identify whether the method would accurately represent both their home range and movement patterns. To assess the suitability of dBBMMs, we compared this novel method with traditional home range estimation methods: minimum convex polygons (MCP) and Kernel density estimators (KDE). Both KDE and MCP incorporated higher levels of Type I and Type II errors, which would lead to biases in our understanding of this species space-use and habitat selection. Although these methods identified some general spatial-temporal patterns, dBBMMs were more efficient at detecting movement corridors and accurately representing long-term shelters sites, showing an improvement over methods traditionally favored in reptile studies. The additional flexibility of the dBBMM approach in providing insight into movement patterns can help further improve conservation and management actions. Additionally, our results suggest that dBBMMs may be more widely applicable in studies that rely on VHF telemetry and not limited to studies employing GPS tags.

Recent literature has reported inaccuracies associated with some popular home range estimators such as kernel density estimation, especially when applied to point patterns of complex shapes. This study explores the use of characteristic hull polygons (CHPs) as a new method of home range estimation. CHPs are special bounding polygons created in GIS that can have concave edges, be composed of disjoint regions, and contain areas of unoccupied space within their interiors. CHPs are created by constructing the Delaunay triangulation of a set of points and then removing a subset of the resulting triangles. Here, CHPs consisting of 95% of the smallest triangles, measured in terms of perimeter, are applied for home range estimation. First, CHPs are applied to simulated animal locational data conforming to five point pattern shapes at three sample sizes. Then, the method is applied to black‐footed albatross (Phoebastria nigripes) locational data for illustration and comparison to other methods. For the simulated data, 95% CHPs produced unbiased home range estimates in terms of size for linear and disjoint point patterns and slight underestimates (8–20%) for perforated, concave, and convex ones. The estimated and known home ranges intersected one another by 72–96%, depending on shape and sample size, suggesting that the method has potential as a home range estimator. Additionally, the CHPs applied to estimate albatross home ranges illustrate how the method produces reasonable estimates for bird species that intensively forage in disjoint habitat patches.

Home ranges provide a conceptual and quantitative representation of animal-habitat associations over time. Methods to estimate home ranges have swiftly progressed by dynamically accounting for various sources of bias. Across that period of growth, one potentially influential source of bias has yet to be robustly scrutinized. Animals inhabiting the terrestrial spatial domain make movement decisions in environments with variable landscape complexity. Despite that reality, home range estimation methods tend to be informed by two-dimensional (2D) data (i.e., x and y coordinates), which analytically presume that these landscapes are flat. This analytical tendency potentially misrepresents the configuration and size of animal home range estimates. To examine the prevalence of this bias, we reviewed literature of terrestrial animal home range estimation published between 2000 and 2019. We recorded the proportion of studies that (1) recognized and (2) incorporated landscape complexity. Over 22.0% (n = 271) of the 1,203 studies recognized the importance of landscape complexity for animal movement. Interestingly, just 0.7% (n = 8) incorporated landscape complexity into the home range estimation. We infer then that landscape complexity represents an important source of bias resulting in the underestimation of terrestrial animal home range size. Given the influence of landscape complexity on terrestrial animal decision making, energetics, and fitness, our analysis highlights an important gap in current home range methodologies. We discuss the implications of our analysis for biased understandings of terrestrial animal spatial ecology with subsequent impacts on management and conservation practices built upon these estimates.

The home range encompasses an animal’s movements as it goes about its normal activity, and several home range estimators have been developed. I evaluated the performance of the Minimum Convex Polygon, Bivariate Normal, and several kernel home range estimators in a geographical information system environment using simulations and a large database of O. pumilio mark-recapture locations. A fixed 90% kernel estimator using Least-Square Cross-Validation (to select the bandwidth) outperformed other methods of estimating home range size and was effective with relatively few capture points. Home range size, core area size, intrasexual overlap, and movement rates among coordinates were higher in female frogs than in male frogs. These measures likely reflect behavioral differences related to territoriality (males only) and parental care (both sexes). The simple Biological Index of Vagility (BIV) generated movement values that scaled well with home range size while revealing more information than home range estimates alone.

The population of Bald Eagles (Haliaeetus leucocephalus) nesting in Louisiana reached precariously low numbers in the early 1970’s, and although the population has since increased markedly, resident eagles remain a species of conservation concern in Louisiana. Using statewide aerial nest survey data from 1975–2008, I documented the history and quantified the recovery of the nesting population in Louisiana. Known active nests increased exponentially from 7 to 387 during this period, exhibiting a mean annual rate of increase of 11.1 ± 0.3% per year with no indications of slowing. By 1990, the nesting population in Louisiana had exceeded all goals of the Southeastern States Bald Eagle Recovery Plan. I used nest location and status data from the most recent year (2007–2008) of the nest monitoring program to examine landscape level nest site selection and success. Success of a nest was not greatly influenced by the physical characteristics around a site, whereas the initial selection of a nest site was most influenced by distance to the historical nest centroid and the proportion of open water and agricultural land within 3 km. Bald Eagles nest during the winter in Louisiana, but they are rarely observed in the state during summer. Therefore, I used satellite GPS transmitters to provide evidence of their undocumented summer migrations and examined parameters such as timing of departure and arrivals, stopover use, routes used, and seasonal and annual variations in these parameters. Marked eagles flew relatively direct routes, exhibiting high route fidelity going between Louisiana and their summering areas, which were spread out across much of Canada and south to Illinois. Using locations from their winter and summering areas, I estimated home range and core area size. Home ranges varied from 19.7–1,997.7 km2, but nesting Bald Eagles had smaller home ranges, on average, than did non-nesting birds. The sex of an individual can play an important role when considering variations in movements, home ranges, and habitat use. Accordingly, I evaluated the published field technique for sex determination of Bald Eagles, and upon documenting misclassifications, I developed a new formula applicable to Louisiana and other southern regions.

Individual studies in wildlife science are indicative rather than conclusive. Although multiple studies can be meta-analyzed in such a way that scientific hypotheses can be tested, robust meta-analyses are often difficult or impossible if variables of interest are not measured in a uniform manner. We hypothesized that measurements, even of basic and unequivocal variables, are rarely standardized in wildlife sciences. We tested this assumption by reviewing randomly selected papers that describe the home range of mammals (n = 25) and birds (n = 25). In these papers, home ranges were calculated using 11 methods and 8 computer programs. The number of radiolocations used to calculate home ranges varied from 9 to >2,000. By estimating home ranges for two radiotelemetry data sets, we demonstrate that home ranges are not comparable if different methods are used and that estimates of home range are not standardized. We assume that measurements of other biological variables are even less consistent across studies. In order to advance wildlife sciences, we believe that standardization initiatives are required at an international level.

ABSTRACTThe African savannah elephant (Loxodonta africana) is listed as endangered on the IUCN Red List. The Ruaha–Rungwa ecosystem supports Tanzania's largest elephant population and remains a critical yet understudied stronghold for elephant conservation in East Africa. Effective conservation in this ecosystem requires understanding elephant movements across large landscapes and varying levels of disturbance. This study addressed the gap by utilizing GPS‐satellite tracking of 28 elephants over a 4‐year period in Tanzania's Ruaha–Rungwa ecosystem. It examined their home ranges and movement patterns inside and outside protected areas and identified key elephant hotspots. The findings aim to improve conservation strategies and mitigate human–elephant conflicts through better habitat management and protection efforts. The study found no significant difference between home range estimates from Minimum Convex Polygon and Adaptive Kernel Density Estimation, with 28 collared elephants averaging 2536 km2 at the 99% isopleth. Home range size varied by age and season but not by sex. Elephants moved faster at night in less protected areas during both wet and dry seasons. Population‐level habitat use was similar across protection levels. Village lands in western Rungwa Game Reserve and its surrounding areas, as well as the eastern regions outside Ruaha National Park, served as key refuge sites during the wet season. Findings suggest elephants in the Ruaha–Rungwa ecosystem require vast areas, with adjacent lands playing a crucial role in their survival. High elephant activity in village lands within the western Rungwa Game Reserve and Lunda‐Mkwambi Game Controlled Area highlights the potential for human–elephant conflict, emphasizing the need to reopen the wildlife corridor for linking Ruaha–Rungwa and Rukwa–Katavi ecosystems. This study offered crucial insights into elephant movement and behavior in a landscape increasingly shaped by human–elephant conflict and habitat fragmentation, informing strategies for connectivity, land‐use planning, and coexistence.

Spatial ecological information is necessary to guide the conservation efforts of river turtles, but it is lacking for many species including the smooth softshell turtle (Apalone mutica). We investigated the spatial ecology of A. mutica in two reaches of the Kaskaskia River in Illinois, USA to determine variables influencing movement rates, the best estimate of home range, and variables influencing home range size. We radio-tracked 28 A. mutica from 2013 to 2014 and used an information theoretic approach to select the best models describing movement and home range estimates. Mean movement rate was 142.3 m/day and was highly variable with some movements >2 km/day. Movement peaked at moderate water temperatures early in the active season, increased at higher water levels for females, and was greater in the higher stream order. The sexes responded differently to environmental variation, with female movement higher in most but not all conditions. The most informative home range estimate was a 95% kernel density estimate using likelihood cross-validation (CVh) smoothing clipped to the river channel. The mean home range size was 18.1 hectares and increased with movement rate, number of radio-locations, and stream order. Most turtles had well-defined home ranges, though a few were possibly nomadic. Our methods also provide a framework for spatial ecological studies of other riverine species.

Understanding the amount of space required by animals to fulfill their biological needs is essential for comprehending their behavior, their ecological role within their community, and for effective conservation planning and resource management. The space-use patterns of habituated primates often are studied by using handheld GPS devices, which provide detailed movement information that can link patterns of ranging and space-use to the behavioral decisions that generate these patterns. However, these data may not accurately represent an animal’s total movements, posing challenges when the desired inference is at the home range scale. To address this problem, we used a 13-year dataset from 11 groups of white-faced capuchins (Cebus capucinus imitator) to examine the impact of sampling elements, such as sample size, regularity, and temporal coverage, on home range estimation accuracy. We found that accurate home range estimation is feasible with relatively small absolute sample sizes and irregular sampling, as long as the data are collected over extended time periods. Also, concentrated sampling can lead to bias and overconfidence due to uncaptured variations in space use and underlying movement behaviors. Sampling protocols relying on handheld GPS for home range estimation are improved by maximizing independent location data distributed across time periods much longer than the target species’ home range crossing timescale.

Providing supplemental food resources for northern bobwhite (Colinus virginanus) has been a common management practice for decades, but its impact on bobwhite home ranges sizes and space use has been debated. Between 2001 and 2007, we established a 397 ha fed study area on Tall Timbers Research Station, which received grain sorghum biweekly along a 19.5 km feed trail. An adjacent 465 ha area was treated as a control with no supplemental food resources. Radio-tagged wild bobwhites were located 3 – 5 times each week throughout the breeding season and we used these locations to calculate home ranges during early breeding season (15 April – 30 June) and late breeding season (1 July – 1 October). We also determined second and third order selection of a 10-m buffer area surrounding the supplemental feed trail using compositional analyses. In total, 552 and 286 bobwhite home ranges were calculated for early and late breeding seasons, respectively. We observed significantly smaller early breeding season home ranges on the fed area (x̄ = 12.3 ha, 95% CL ± 0.6) relative to the control (x̄ = 17.4 ha ± 1.9). Average length of feed trail within home ranges was 583 m and 710 m for early and late breeding season home ranges, respectively. The feed trail buffer area was 3 to 5% of all home ranges. Home range placement was not random with high selection preference for the feed trail buffer area. However, there was a low selection preference for the feed trail buffer area within home ranges. Average distances to the feed trail for bobwhite locations (x̄ = 64.4 m ± 16.1) was similar to random locations (x̄ = 74.4 m ± 16.9). The effect of supplemental feeding on bobwhite home ranges size and resource use was greatest during the early breeding season when food and useable habitat were likely lower in availability. On our study area, supplemental food resources distributed along a feed trail impacted bobwhites during a critical transition period from the spring prescribed burning season to the early breeding season.

The small Indian mongoose (Urva auropunctata) is the primary terrestrial wildlife rabies reservoir on at least four Caribbean islands, including Puerto Rico. In Puerto Rico, mongooses represent a risk to public health, based on direct human exposure and indirectly through the transmission of rabies virus to domestic animals. To date, the fundamental ecological relationships of space use among mongooses and between mongooses and domestic animals remain poorly understood. This study is the first to report mongoose home range estimates based on GPS telemetry, as well as concurrent space use among mongooses and free roaming domestic dogs (FRDD; Canis lupus familiaris). Mean (± SE) home range estimates from 19 mongooses in this study (145 ± 21 ha and 60 ± 14 ha for males and females, respectively) were greater than those reported in prior radiotelemetry studies in Puerto Rico. At the scale of their home range, mongooses preferentially used dry forest and shrubland areas, but tended to avoid brackish water vegetation, salt marshes, barren lands and developed areas. Home ranges from five FRDDs were highly variable in size (range 13–285 ha) and may be influenced by availability of reliable anthropogenic resources. Mongooses displayed high home range overlap (general overlap index, GOI = 82%). Home range overlap among mongooses and FRDDs was intermediate (GOI = 50%) and greater than home range overlap by FRDDs (GOI = 10%). Our results provide evidence that space use by both species presents opportunities for interspecific interaction and contact and suggests that human provisioning of dogs may play a role in limiting interactions between stray dogs and mongooses.

To investigate the roaming of Indigenous community dogs and potential interaction with wild dogs and dingoes. Cross-sectional survey and longitudinal follow-up study. Six remote Indigenous communities in Cape York Peninsula and Arnhem Land in northern Australia were selected. Hair samples were collected from community dogs and microsatellite DNA analyses were used to determine hybrid (>10% dingo DNA) status. Dogs were fitted with GPS collars and home range (ha) was estimated during monitoring periods of up to 3 days. In Cape York Peninsula, 6% of the 35 dogs sampled were dingo hybrids, whereas in Arnhem Land 41% of the 29 dogs sampled were hybrids. The median extended home range was estimated to be 4.54 ha (interquartile range, 3.40 - 7.71). Seven community dogs were identified with an estimated home range > 20 ha and home ranges included the bushland surrounding communities. No significant difference in home ranges was detected between hybrid and non-hybrid dogs. Study results provide some evidence (dingo hybridisation, bushland forays) of the potential interaction between domestic and wild dogs in northern Australia. The nature of this interaction needs further investigation to determine its role in disease transmission; for example, in the case of a rabies incursion in this region.