Effects of Point Pattern Shape on Home-Range Estimates

Abstract: Home-range estimators are commonly tested with simulated animal locational data in the laboratory before the estimators are used in practice. Although kernel density estimation (KDE) has performed well as a home-range estimator for simulated data, several recent studies have reported its poor performance when used with data collected in the field. This difference may be because KDE and other home-range estimators are generally tested with simulated point locations that follow known statistical distributions, such as bivariate normal mixtures, which may not represent well the space-use patterns of all wildlife species. We used simulated animal locational data of 5 point pattern shapes that represent a range of wildlife utilization distributions to test 4 methods of home-range estimation: 1) KDE with reference bandwidths, 2) KDE with least-squares cross-validation, 3) KDE with plug-in bandwidths, and 4) minimum convex polygon (MCP). For the point patterns we simulated, MCP tended to produce more accurate area estimates than KDE methods. However, MCP estimates were markedly unstable, with bias varying widely with both sample size and point pattern shape. The KDE methods performed best for concave distributions, which are similar to bivariate normal mixtures, but still overestimated home ranges by about 40–50% even in the best cases. For convex, linear, perforated, and disjoint point patterns, KDE methods overestimated home-range sizes by 50–300%, depending on sample size and method of bandwidth selection. These results indicate that KDE does not produce home-range estimates that are as accurate as the literature suggests, and we recommend exploring other techniques of home-range estimation.     

Evaluation of Root-n Bandwidth Selectors for Kernel Density Estimation

ABSTRACT The kernel density estimator is used commonly for estimating animal utilization distributions from location data. This technique requires estimation of a bandwidth, for which ecologists often use least-squares cross-validation (LSCV). However, LSCV has large variance and a tendency to under-smooth data, and it fails to generate a bandwidth estimate in some situations. We compared performance of 2 new bandwidth estimators (root-n) versus that of LSCV using simulated data and location data from sharp-shinned hawks (Accipter striatus) and red wolves (Canis rufus). With simulated data containing no repeat locations, LSCV often produced a better fit between estimated and true utilization distributions than did root-n estimators on a case-by-case basis. On average, LSCV also provided lower positive relative error in home-range areas with small sample sizes of simulated data. However, root-n estimators tended to produce a better fit than LSCV on average because of extremely poor estimates generated on occasion by LSCV. Furthermore, the relative performance of LSCV decreased substantially as the number of repeat locations in the data increased. Root-n estimators also generally provided a better fit between utilization distributions generated from subsamples of hawk data and the local densities of locations from the full data sets. Least-squares cross-validation generated more unrealistically disjointed estimates of home ranges using real location data from red wolf packs. Most importantly, LSCV failed to generate home-range estimates for >20% of red wolf packs due to presence of repeat locations. We conclude that root-n estimators are superior to LSCV for larger data sets with repeat locations or other extreme clumping of data. In contrast, LSCV may be superior where the primary interest is in generating animal home ranges (rather than the utilization distribution) and data sets are small with limited clumping of locations.     

LANDSCAPE PROPORTIONS VERSUS MONTE CARLO SIMULATED HOME RANGES FOR ESTIMATING HABITAT AVAILABILITY

Abstract: Wildlife researchers often test whether animals use resources disproportionately relative to availability (i.e., selectively). However, the traditional estimate of availability at the landscape scale (resource proportions on the landscape) may be inaccurate and lead to false conclusions. We calculated the chance of falsely finding selection (type I error rate) when the traditional estimate of availability is used. True availability was estimated by Monte Carlo simulations with randomly located home ranges and compared to the traditional estimate to calculate type I error rates. Tests were conducted with α = 0.05 for different home-range sizes (1 to 1,000 km²) and 4 habitat patterns. Landscape proportions did not equal proportions of habitats in random home ranges (traditional estimate ≠ true availability). Type I error rates were ≥0.24 and increased with number of animals tested and decreased with home-range size and number of habitats. Therefore, researchers should use randomly located home ranges instead of landscape proportions to estimate availability at the landscape scale. We evaluated a goodness-of-fit test for comparing habitat proportions between randomly located home ranges and observed home ranges. Type I error rates for this method were ≤0.08, regardless of number of animals, home-range size, and number of habitats tested. We evaluated this method for 2 species with different home-range sizes and predicted habitat selection patterns: mountain lions (Puma concolor, ∼ 700 km², relatively nonselective) and mule deer (Odocoileus hemionus, ∼ 16 km², relatively selective). This method yielded results consistent with predictions, whereas the traditional method using landscape proportions to estimate availability did not. Randomly located, simulated home ranges are superior to landscape proportions for estimating availability.     

The home-range concept: are traditional estimators still relevant with modern telemetry technology?

Recent advances in animal tracking and telemetry technology have allowed the collection of location data at an ever-increasing rate and accuracy, and these advances have been accompanied by the development of new methods of data analysis for portraying space use, home ranges and utilization distributions. New statistical approaches include data-intensive techniques such as kriging and nonlinear generalized regression models for habitat use. In addition, mechanistic home-range models, derived from models of animal movement behaviour, promise to offer new insights into how home ranges emerge as the result of specific patterns of movements by individuals in response to their environment. Traditional methods such as kernel density estimators are likely to remain popular because of their ease of use. Large datasets make it possible to apply these methods over relatively short periods of time such as weeks or months, and these estimates may be analysed using mixed effects models, offering another approach to studying temporal variation in space-use patterns. Although new technologies open new avenues in ecological research, our knowledge of why animals use space in the ways we observe will only advance by researchers using these new technologies and asking new and innovative questions about the empirical patterns they observe.     

Use of space and habitat selection by roe deer Capreolus capreolus in a Mediterranean coastal area: how does woods landscape affect home range?

The home-range dynamics and habitat selection of nine roe deer were studied from March 1994 to August 1994 in the Maremma Natural Park along the Tyrrhenian coast of Italy. The habitat was highly fragmented, with open agricultural fields prevailing in the study area (57%); the climate was Mediterranean. Data on spatial behaviour were collected by radio-tracking techniques. Habitat selection and structure were investigated by compositional and landscape analysis, both within the study area and within the home ranges. Animals of our sample showed spatial-use patterns varying from stationary to roaming. Stationary individuals used small home ranges while roaming ones moved, especially during the reproductive period in July and August. The percentage and structure of woodlands influenced the size of home ranges and the behaviour of males: stationary males used large amounts of woodlands within their home ranges and showed a territorial behaviour whereas males that used a high percentage of fields showed wider home ranges even during the territorial period. Females seemed to be less influenced by the presence and patch-structure of woodland within their home range. Landscape structure and habitat composition seemed to be important factors influencing the spatial behaviour of this roe deer population.     

Socio-spatial ecology of pine marten (Martes martes) in conifer forests,Ireland

Understanding the social organisation and spacing patterns of wildlife populations is an important aspect of conservation management and applied science. The present study investigated the spatial ecology of pine marten (Martes martes) inhabiting conifer forests in Ireland, the largest habitat resource available for the species. It represented the first study of pine marten spacing patterns in Irish conifer forests. Pine marten (n?=?7; five males and two females) were live-trapped and radio-tracked for between 4 and 10 months from March 2008 to March 2009. Mean annual home-range estimates (95 % fixed kernel) for males (150.7 ha) were generally larger than those of females (90.2 ha). There was considerable inter-seasonal overlap in home ranges (approx 85 %) with less inter-sexual (12.0 %) or intra-sexual (11.8 %) overlap, although the sample size of individuals for comparison was small. Pine marten home ranges were stable from season to season. Core ranges varied in size from 10.6 to 104.1 ha, and as a mean percentage of home-range area were 22.9 and 42.5 % for males and females, respectively. In terms of forest management, potential under occupancy of available space by pine marten and vulnerability of very small core ranges to clear felling practices needs further research to determine any impacts on individuals and populations.     

Home range and habitat use by the serotine bat, Eptesicus serotinus, in England

Serotine nursery roosts with less than 20 bats were found to have home ranges of at least 24 to 77 km² and core areas of activity from 13 to 33 km². The size of the range may have increased further if more individuals had been tracked, as three of the four colonies studied had not reached their asymptotes. The total home-range area covered by four serotine colonies was 127.36 km². Excluding non-breeding bats, a density of one bat per 120 ha was estimated. However, actual density was likely to be higher if there were additional non-breeding females and immatures that were not in nursery roosts. Colonial home ranges and core areas overlapped, with individuals from different colonies feeding at the same sites. Individual home ranges ( n = 32) varied from 0.16 to 47.58 km², but these were not used exclusively by one individual. Around the colonial core area and breeding roosts, home ranges were used by all individuals from a single colony. It is only further from the core area that ranges appeared to be used by individuals. The distance from roost to feeding areas varied by up to 7.4 km, but the bat usually commuted along lines of trees and hedges and over pastures. This resulted in greater distances being travelled than if they had flown by a direct route. On average, individuals commuted distances of 8 km each night between feeding areas, with a maximum distance of over 41 km. They visited between 0 and 10 feeding sites each night (mean = 2.89).     

蜂桶寨自然保护区小熊猫巢域初步研究

2002年5~11月,在蜂桶寨自然保护区利用无线电遥测技术对6只小熊猫的巢域利用进行了初步研究。结果表明,6只戴颈圈个体M1、M2、M3、F1、F2、F3的巢域面积分别为330·26hm~2、135·18hm~2、190·67hm~2、98·23hm~2、141·60hm~2、204·80hm~2;雄性个体平均巢域面积为218·70hm~2,雌性个体为148·21hm~2。小熊猫个体间巢域重叠普遍,平均重叠率达25·33%,其中雄性个体之间为26·00%,雌性个体之间为23·67%,两性个体之间为25·67%。可能受人为干扰的影响,M1在6只监测个体中巢域面积、日均移动距离均为最大。     

Home ranges in adult Scandinavian brown bears (Ursus arctos): effect of mass, sex, reproductive category, population density and habitat type

Annual home-range size indices for 36 male and 52 female adult brown bears Ursus arctos in two study areas in central and northern Scandinavia were estimated to evaluate factors believed to influence home-range size. Male home ranges were larger than home ranges of lone females after controlling for the sexual size dimorphism acting on metabolic needs. Further, home ranges of females with cubs were smaller than home ranges of lone females and females with yearlings. Thus, differences in metabolic need were not able to explain the variation in range size among females of different reproductive categories or between males and females, suggesting roaming behaviour of males in this promiscuous species. Home-range size in both males and females was inversely related to population density along a density gradient that was not linked to food availability. This contradicts the hypothesis that females use the minimum areas that sustain their energy requirements. However, on a large geographical scale a negative relationship between range size and food availability was evident. The annual home ranges in inland boreal environments in Scandinavia are the largest reported for brown bears in Eurasia, and similar to those in inland boreal and montane environments in North America.     

Estimating utilization distributions with kernel versus local convex hull methods

Estimates of utilization distributions (UDs) are used in analyses of home-range area, habitat and resource selection, and social interactions. We simulated data from 12 parent UDs, representing 3 series of increasingly intense space-use patterns (clustering of points around a home site, restriction of locations to a network of nodes and corridors, and dominance of a central hole in the UD) and compared the ability of kernel density estimation (KDE) and local convex hull (LCH) construction to reconstruct known UDs from samples of 10, 50, 250, and 1,000 location points. For KDE, we considered 4 bandwidth selectors: the reference bandwidth, least-squares cross-validation (LSCV), direct plug-in (DPI), and solve-the-equation (STE). For the sample sizes and UD patterns tested here, KDE achieved significantly higher volume-of-intersection (VI) scores with known parent UDs than did LCH; KDE also provided less biased home-range area estimates under many conditions. However, LCH minimized the UD volume that occurred outside the true home range boundary (V_out). Among the KDE bandwidth estimators, relative performance depended on the type and intensity of space use patterns, sample size, and the metric used to evaluate performance. Biologists should use KDE for UD and home range estimation within a probabilistic context, unless their objective is to exclude potentially unused areas by defining the area delimited by data. © 2011 The Wildlife Society.     

A comparison of trapping- and radiotelemetry-based estimates of home range of the neotropical opossum Philander frenatus

Home ranges of individuals of the gray four-eyed opossum Philander frenatus were studied by capture–mark–recapture (CMR) and radiotelemetry, within a set of eight Atlantic Forest fragments surrounded by a grassland matrix in the state of Rio de Janeiro, southeastern Brazil. Trapping sessions were carried out in all the forest fragments and in the grassland matrix. Adult individuals were fitted with radio-collar transmitters and monitored throughout the night. Locations were obtained by the “homing-in on the animal” method. Home-range sizes of the individuals with five or more captures or locations were estimated through the minimum convex polygon method. Home-range sizes estimated by radiotelemetry ranged from 0.6 to 7.4 ha (n=8), and by CMR ranged from 0.1 to 12.1 ha (n=17); home-range sizes estimated by radiotelemetry did not differ significantly from those based on CMR. However, comparing radiotelemetry with different CMR designs, the former estimates were larger than those based on either CMR using a single grid or CMR using two grids, but not larger than those based on multiple-grid CMR. In specific cases, animals monitored via radiotelemetry for only one or two nights showed larger home ranges than most individuals for which home ranges were estimated by CMR. Two individuals for which home-range sizes were estimated by both techniques showed larger home ranges when data from radiotelemetry were used. These data indicated that CMR can provide results comparable to radiotelemetry when multiple grids, spread across the landscape, are used, although this necessitates an intensive trapping effort. On the other hand, single- and double-grid CMR tend to underestimate home-ranges compared to radiotelemetry.     

A Critical Review of Home Range Studies

Abstract No consensus currently exists for the methods of estimation of home range size or for reporting home-range analysis results. Studies currently employ a variety of disparate methods or provide inadequate information for reproducing their analyses. We reviewed 161 home range studies published in 2004, 2005, and 2006 to assess what methods are currently employed and how results are reported. We found that home range reporting was generally inadequate for reproducing studies; that the methods employed varied considerably; that home range estimates were often reported and analyzed using inappropriate methods; and that many comparisons were made between studies that may produce spurious results. We urge for minimum editorial standards for reporting home range studies and we urge researchers to follow a unified methodology for estimating animal home ranges. We supply recommendations for such reporting and methodology. These recommendations will increase the reproducibility of studies and allow for more robust comparisons between studies.     

Dispersal of Yearling Pronghorns in Western South Dakota

Abstract: We captured and radiocollared 57 pronghorn (Antilocapra americana) fawns in western South Dakota, USA, during May 2002–2003 and radiotracked them through 15 months of age, by which time all surviving individuals had established a permanent home range. We classified 56% (n = 19) of fawns as dispersers and 44% (n = 15) as residents. Eighty-four percent (n = 16) of dispersers departed natal home ranges in late October and occupied winter home ranges for 102–209 days before dispersing to permanent home ranges during April 2003 and 2004. Dispersal distances from natal ranges to permanent home ranges varied from 6.2–267.0 km. Winter home-range sizes for all individual pronghorns varied from 39.4–509.6 km. Permanent home-range size for all individuals varied from 15.5–166.1 km². Mean 95% permanent home-range size differed (P = 0.06) between residents (x̄ = 97.3 ± 15.1 km²) and dispersers (x̄ = 48.6 ± 16.0 km²), but was similar (P = 0.97) among sexes. Mean dispersal distance from natal to permanent home ranges was similar (P = 0.35) for males (x̄ = 54.2 ± 21.0 km) and females (x̄ = 26.3 ± 19.9 km). We suggest that habitat quality (i.e., patchiness) and pronghorn density, in part, stimulated dispersal. We hypothesize that as habitat patch size decreases, home range sizes and distance traveled during predispersal and dispersal movements by pronghorns will increase.     

Seasonal locomotion and home-range characteristics of European hares (<Emphasis Type="Italic">Lepus europaeus</Emphasis>) in an arable region in central Germany

We studied seasonal use of space by 38 radio-tracked European hares in an arable region in central Germany over 5 years. Mean distance between successive daytime and nighttime fixes of a hare amounted to 226 m, and an average distance of 172 m was recorded for successive daytime fixes. The hares shifted the centres of their home ranges from one 2-month period to the next by an average distance of 131 m and over a time span of 11.2 months by 216 m. The size of their 2-month home ranges (MCP 95) averaged 21 ha; the variation between individuals was high. Night ranges were larger than day ranges. Mean size of seasonal home ranges—based on daytime and nighttime fixes—remained largely constant throughout the year. Home-range size increased as the day-to-night distance increased, as the number of used habitat elements increased and as frequency of use of inner field parts during the daytime increased. Home-range size was inversely related to population density. On average, 32% of the area of the home range of a hare overlapped with the home range of a neighbouring conspecific. When related to the population density of adult hares in spring, the home range of a single hare was estimated to overlap with 13–21 home ranges of adults as well as an indefinite number of juveniles. The large intraseasonal variation in locomotion and home-range characteristics is interpreted in relationship to the social structure of European hares.     

Estimation of the maximum utilization area including home range and peripheral sites

There is increasing evidence that occasional utilization area (peripheral sites), in addition to typical utilization area (home range), is important for wildlife conservation and management. Here we estimated the maximum utilization area (MUA), including both typical and occasional utilization areas, based on asymptotic curves of utilization area plotted against sample size. In previous studies, these curves have conventionally been plots of cumulative utilization area versus sample size, but this cumulative method is sensitive to stochastic effects. We propose a new method based on simulation studies where outcomes of replicated simulations are averaged to reduce stochastic effects. In this averaged method, possible combinations of sample size with the same number of location data replicated from a dataset were averaged and applied to the curves of utilization area. The cumulative method resulted in a large variation of MUA estimates, depending on the start date as well as total sample size of the dataset. In the averaged method, MUA estimates were robust against changes in the start date and total sample size. The large variation of MUA estimates arose because location data on any day including the start date are affected by unpredictable effects associated with animal activity and environmental conditions. In the averaged method, replicates of sample size resulted in a reduction of temporal stochasticity, suggesting that the method stably provides reliable estimates for MUA.     

啮齿动物的巢区面积估算法

巢区(Home range)是动物在其巢附近进行取食、生殖、育幼等日常活动的区域(Burt 1940)。标志流放法是应用最广的调查啮齿动物巢区的方法,尤其是按方格式布笼。但对同一野外调查结果,由于估算方法不同,巢区估算值相差很大,并且至今尚无学者提出一致公认的估算法。1980年5-10月,我们在青海省门源县的高寒草甸生态系统定位站调查根田鼠的巢区,按多种估算法对同一批实际调查结果估计了巢区面积,并对结果进行了分析比较,检查其特点和优缺点,并提出修正平均值法,作为我们今后讨论根田鼠巢区动态的基础。 对方格式布笼的调查结果进行巢区的估算方法,基本上分二大类,图形法和概率性模型法。图形法是按照捕点分布划出巢区,并直接求出巢区面积,如最小面积法,包括或不包括周边地带法,最大距离法和复合散布图法。     

Territorial behavior of the plethodontid salamander Plethodon kentucki: influence of habitat structure and population density

To investigate the possible influence of variation in ecological and demographic factors on the spatial organization of the terrestrial plethodontid salamander Plethodon kentucki, I conducted a 3-year capture-recapture study and determined home-range characteristics and spatial relationships of individuals at two field sites that differed in predominant cover type and population density. Home ranges of adults were fixed and the home ranges of same-sex adult neighbors were mostly exclusive. The spatial arrangement of adult home ranges exhibited overall regularity or regularity within aggregations, whereas the distribution of juvenile home ranges was usually random. Analysis of nearest-neighbor sex indicated a positive intersexual association of adult home ranges. Removal studies provided evidence for defense of adult home ranges only at the high-density site. The distribution of home ranges was influenced by the presence of cover objects, but there was no significant relationship between adult body size and percent of home-range area with cover. Males overlapped the home ranges of gravid females significantly more often than those of non-gravid females, indicating that the distribution of gravid females had a strong influence on the distribution of male home ranges. In laboratory tests, increased male-male aggression during the breeding season suggests that males may compete for access to mates. At the high-density site, larger males may have benefited by having greater reproductive success than smaller males because they were more dominant and their home ranges overlapped a greater number of gravid-female home ranges. My results indicate that habitat structure and population density may influence the spatial organization and mating system of P. kentucki. Received: 26 May 1997 / Accepted: 9 October 1997     

Population ecology of the white-nosed coati (Nasua narica) on Barro Colorado Island, Panama

The white-nosed coati, Nasua narica , is a common Neotropical carnivore with a social structure of band-living adult females and solitary adult males. A coati population on Barro Colorado Island, Panama, was studied over a four-year period by mark-recapture, radiotelemetry. and direct observation of habituated individuals. The population density was approximately 51.5 individuals/km² and the sex ratio was 1:1. Band size varied from six to 26 individuals (mean = 15.3) with extensive fluctuation within and between years. Mean foraging group size was smaller (7.2 individuals) than population group size, and fluctuated with food availability, synchronous parturition, and the emigration of mature males. Mean home-range size of six bands was 0.33 km², and ranges of adjacent bands overlapped from 0–66%. One band fissioned during the study; however, the resulting bands did not disperse from the original home range. Seven adult males had a mean home-range size of 0.37 km², each extensively overlapping the home ranges of several other males. Observations of 10 adult males whose natal bands were known indicate that when males disperse they do not simultaneously leave the band's home range. Rather, their home ranges remain within or broadly overlapping those of their natal bands. This dispersal pattern is unusual within the order Carnivora.     

Memory keeps you at home: a mechanistic model for home range emergence

Despite its central place in animal ecology no general mechanistic movement model with an emergent home-range pattern has yet been proposed. Random walk models, which are commonly used to model animal movement, show diffusion instead of a bounded home range and therefore require special modifications. Current approaches for mechanistic modeling of home ranges apply only to a limited set of taxa, namely territorial animals and/or central place foragers. In this paper we present a more general mechanistic movement model based on a biased correlated random walk, which shows the potential for home-range behavior. The model is based on an animal tracking a dynamic resource landscape, using a biologically plausible two-part memory system, i.e. a reference- and a working-memory. Our results show that by adding these memory processes the random walker produces home-range behavior as it gains experience, which also leads to more efficient resource use. Interestingly, home-range patterns, which we assessed based on home-range overlap and increase in area covered with time, require the combined action of both memory components to emerge. Our model has the potential to predict home-range size and can be used for comparative analysis of the mechanisms shaping home-range patterns.     

两种核域估算方法在野生藏狐家域研究中的比较

核域是野生动物重点利用、含有更多居所、隐蔽场所及依赖性食物资源的区域。关于核域的确定和划分一直存在争议,且很难找到客观的并能准确展示动物生物学信息的数理统计方法。2006 年4 月,2006 年9 ～10月和2007 年1 ～4 月,我们共记录青海省都兰县沟里乡3 只藏狐的199 个活动位点,通过固定核空间(Fixed kernel)和调和平均值(Harmonic mean)模型,结合独立区域法的计算原理,估算藏狐的核域,并对两种估算方法进行比较分析,以确定更理想的核域数学模型。研究结果表明:(1)两种估算方法中核域面积均受到家域总面积的影响,固定核空间法的核域面积受家域总面积影响较大,而且变量系数不稳定; (2) 随着样本量的增加,固定核空间法估算的核域面积逐渐减小,而调和平均值法则增大,前者从数理统计上更好地描述了藏狐对家域资源的利用分布,后者更符合客观实际;(3)调和平均值法在划定家域边界时容易包括不属于家域范围的区域,而在估算核域时可以在一定程度上克服该缺陷;(4)调和平均值法能够真实地反应动物的活动中心,估算的核域中包含有更多的活动位点。因此,尽管在估算藏狐核域时固定核空间法有稳定的计算结果等优点,调和平均值法为更理想的核域估算模型。