A Mechanistic Approach to Evaluation of Umbrella Species as Conservation Surrogates

Abstract:  Although species with large area requirements are sometimes used as umbrella species, their general utility as conservation tools is uncertain. We surveyed the species diversity of birds, butterflies, carabids, and forest-floor plants in forest sites across an area (1600 km²) in which we delineated large breeding home ranges of Northern Goshawk ( Accipiter gentilis ). We tested whether protection of the home ranges could serve as an effective umbrella to protect sympatric species of the four taxa. We also used an empirical habitat model of occupancy of home range to examine mechanisms by which the Northern Goshawk acts as an umbrella species. Among species richness, abundance, and species composition of the four taxa, only abundance and species composition of birds differed between sites located inside and outside home ranges, which was due to greater abundance of bird species that were prey of Northern Goshawks inside the home ranges. Thus, although home range indicated areas with high abundance of certain bird prey species, it was not effective as an indicator of the species diversity of all four taxa. We also did not find any difference in species richness, abundance, and species composition between sites predicted as occupied and unoccupied using the habitat model. In contrast, when we selected sites on the basis of each habitat variable in the model, habitat variables that selected sites either in agricultural or forested landscapes encompassed sites with high species richness or particular species composition. This result suggests that the low performance of the Northern Goshawk as an umbrella species is due to this species' preference for habitat in both agricultural and forested landscapes. Species that can adjust to changes in habitat conditions may not act as effective umbrella species despite having large home ranges.     

Neighboring groups and habitat edges modulate range use in Phayre’s leaf monkeys (<Emphasis Type="Italic">Trachypithecus phayrei crepusculus</Emphasis>)

An animal’s use of space may be strongly influenced by habitat edges and neighboring conspecifics encountered in and around its home range. Habitat edges are known to affect species density and distribution, but their impact on home range use is largely unknown. Additionally, among large animals, interactions with neighbors become particularly important as increasing home range size leads to decreasing exclusivity of resource use, but the effect of neighbors on home range use remains poorly understood. Here, we examine the influence of neighbors and habitat edges on the ranging patterns of three groups of Phayre’s leaf monkeys (Trachypithecus phayrei crepusculus) in northeast Thailand over a period of more than 2 years. The study animals occupied dry evergreen forest, and adjacent patches of dry dipterocarp forest created a habitat edge and formed barriers between some groups. We found that the use of home range interiors was 50–90% higher than the border areas, indicating concentrated use of resources within the home range. The use of peripheral areas was influenced by social organization, the presence of neighboring groups, and forest edges. While one multimale group showed no particular habitat preference, two single-male groups preferred areas bordering dry dipterocarp habitat and avoided areas bordering neighboring groups, suggesting that the threat of neighbors mediated border presence. Additionally, groups may have been attracted to the forest edge, where conspecific competitors are absent and increased sunlight may increase resource abundance and/or quality. This study revealed that the use of border areas can be modulated by neighboring groups and habitat edges, thereby adding to our understanding of home range use among territorial species in heterogeneous habitats.     

Kernel-based home range method for data with irregular sampling intervals

Studies of habitat selection and movements often use radio-tracking data for defining animal home ranges. Home ranges (HR) can be approximated by a utilization density distribution (UD) that instead of assuming uniform use of areas within HR boundary provides a probabilistic measure of animal space use. In reality, radio-tracking data contain periods of frequent autocorrelated observations interspersed with temporally more independent observations. Using such temporally irregular data directly may result in biased UD estimates, because areas that have been sampled intensively receive too much weight. The problem of autocorrelation has been tackled by resampling data with an appropriate time interval. However, resampling may cause a large reduction in the data set size along with a loss of information. Evidently, biased UD estimates or reduction in data may prejudice the results on animal habitat selection and movement. We introduce a new method for estimating UDs with temporally irregular data. The proposed method, called the time kernel, accounts for temporal aggregation of observations and gives less weight to temporally autocorrelated observations. A further extension of the method accounts also for spatially aggregated observations with relatively low weights given to observations that are both temporally and spatially aggregated. We test the behaviour of the time kernel method and its spatiotemporal version using simulated data. In addition, the method is applied to a data set of brown bear locations.     

Iberian Lynx in a Fragmented Landscape: Predispersal, Dispersal, and Postdispersal Habitats

Abstract: Applied conservation biology must provide solutions for the conservation of species in modern landscapes, where prime habitats are being continuously fragmented and altered and animals are restricted to small, nonviable populations. We studied habitat selection in a fragmented population of endangered Iberian lynx (   Lynx pardinus ) by examining 14 years of radiotracking data obtained from lynx trapped in two different source areas. Habitat selection was studied independently for predispersal lynx in the source areas, for dispersing individuals through the region, and for postdispersing animals, most of which settled far from their point of origin. A multivariate analysis of variance showed that habitat use differed significantly among these phases and between area of origin, but not between sexes. The habitat type most used, and best represented within home ranges, was the mediterranean scrubland. Pine plantations were also important during and after dispersal. The rest of the habitats were either avoided (open habitats) or used according to availability ( pine and eucalyptus plantations) by dispersing lynx. Differences due to lynx origin were detected only during predispersal and dispersal and were observed because animals from each area had different habitat availability. Lynx with established territories did not use areas at random. They occupied patches of mediterranean scrubland more often than would be expected from scrubland availability during predispersal; the rest of the habitats were included within home ranges less than would be expected from their availability in the landscape. Results indicate that dispersing animals may use habitats of lower quality than habitats used by resident individuals, which suggests that conservation strategies applied across regions might be a viable objective.     

Understanding determinants of home range behaviour of feral cats as introduced apex predators in insular ecosystems: a spatial approach

The introduced feral cat (Felis catus) is a widespread generalist with flexible social behaviour and an apex predator without major interspecific competitors in insular ecosystems that evolved in the absence of predators. Mechanistic definitions consider an animal’s home range to be the spatial expression of a cognitive map that is kept up-to-date with the status of critical resources that contribute to animal fitness. We assumed there are two major determinants structuring the home range of cats as apex predators in insular ecosystems: the distribution of critical food resources and conspecific distribution. We hypothesized that cats structure their home ranges by optimizing the use of staple critical food resources and that as a consequence of the presence of rich resources cats tend to socialise, aggregate and share space. We carried out spatial analyses using location data for feral cats tracked using lightweight GPS collars in conjunction with the suitability value of rabbit patches and their associated ownership costs for cats within a New Zealand braided-river environment. Cat home ranges and spatial distribution, especially for females, were related to the inclusion of rabbit patches within home ranges with higher mean value than the average of neighbourhood patches in the landscape. Cats showed solitary behaviour but tolerance to conspecific presence by sharing high-use areas and high-value rabbit patches, mostly at different times, resulting in occasional encounters among males and females. Home range size and patterns of spatial overlap were dependant on sex and season. Solitary spacing patterns as consequence of innate preferences together with resource constraints may regulate feral cat population densities.     

Spatiotemporal dynamics of habitat use by koalas: the checkerboard model

Animal movement patterns and use of space depend upon food and nonfood resources, as well as conspecific and heterospecific interactions, but models of habitat use often neglect to examine multiple factors and rarely include marsupials. We studied habitat use in an Australian population of koalas (Phascolarctos cinereus) over a 6-year period in order to determine how koalas navigate their environment and partition limited patchy food and nonfood resources. Tree selection among koalas appears to be mediated by folar chemistry, but nonfood tree selection exerts a major impact on home range use due to thermoregulatory constraints. Koalas moved on a daily basis, during both day and night, but daytime resting site was not necessarily in the same location as nighttime feeding site. Koalas had substantial home range overlap in the near absence of resource sharing with less than 1% of trees located in areas of overlap used by multiple koalas. We suggest that koala spatiotemporal distribution and habitat use are probably based upon a community structure of individuals, with a checkerboard model best describing overlap in home range area but not in resource use. Nonfood refugia and social networks should be incorporated into models of animal range and habitat use.     

Landscape evaluation in conservation: molecular sampling and habitat modeling for the Iberian lynx.

Conservation of endangered species requires comprehensive understanding of their distribution and habitat requirements, in order to implement better management strategies. Unfortunately, this understanding is often difficult to gather at the short term required by rapidly declining populations of many rare vertebrates. We present a spatial habitat modeling approach that integrates a molecular technique for species detection with landscape information to assess habitat requirements of a critically endangered mammalian carnivore, the Iberian lynx (Lynx pardinus), in a poorly known population in Spain. We formulated a set of model hypotheses for habitat selection at the spatial scale of home ranges, based on previous information on lynx requirements of space, vegetation, and prey. To obtain the required data for model selection, we designed a sampling protocol based on surveys of feces and their molecular analysis for species identification. After comparing candidate models, we selected a parsimonious one that allowed (1) reliable assessment of lynx habitat requirements at the scale of home ranges, (2) prediction of lynx distribution and potential population size, and (3) identification of landscape management priorities for habitat conservation. This model predicted that the species was more likely to occur in landscapes with a higher percentage of rocky areas and higher cover of bushes typical of mature mediterranean shrubland mosaics. Its accuracy for discriminating lynx presence was approximately 85%, indicating high predictive performance. Mapping model predictions showed that only 16% of the studied areas constitute potential habitat for lynx, even though the region is dominated by large extents of well-preserved native vegetation with low human interference. Habitat was mostly clumped in two nearby patches connected by vegetation adequate for lynx dispersal and had a capacity for 28-62 potential breeding territories. The lynx population in Sierra Morena is probably the largest persisting today, but it is still critically small for optimism about its long-term persistence. Model results suggest habitat conservation and restoration actions needed for preserving the species, including reconciliation of hunting management with preservation of mature shrubland over large areas (particularly in rocky landscapes). The approach presented here can be applied to many other species for which the ecological information needed to develop sound habitat conservation strategies is lacking.     

Managing Boreal Forest Landscapes for Flying Squirrels

Abstract: Flying squirrel (Pteromys volans) populations have declined severely during the past few decades, and the species has become a focal species in forest management and the conservation debate in Finland. We compared landscape structure around known flying squirrel home ranges with randomly chosen forest sites to determine which landscape patterns characterize the areas occupied by the species in northern Finland. We sought to identify the key characteristics of the landscape that support the remaining flying squirrel populations. We analyzed landscape structure within circular areas with 1- and 3-km radii around 63 forest sites occupied by flying squirrels, and around 96 random sites. We applied stepwise analysis of the landscape structure where landscapes were built up step-by-step by adding patch types in order of their suitability for the flying squirrel. The land-use and forest-resource data for the analysis were derived from multisource national forest inventory and imported to a geographical information system. Landscape patch types were divided into three suitability categories: breeding habitat (mixed spruce-deciduous forests); dispersal habitat ( pine and young forests); and unsuitable habitat ( young sapling stands, open habitats, water). Flying squirrel landscapes contained more suitable breeding habitat patches and were better connected by dispersal habitats than random landscapes. Our results suggest that for the persistence of the flying squirrel, forest managers should 1) maintain a deciduous mixture, particularly in spruce-dominated forests; 2) maintain physical connectivity between optimal breeding habitats; and 3) impose coarse-grained structures on northeastern Finnish landscapes at current levels of habitat availability.     

Conservation of Fragmented Populations

In this paper we argue that landscape spatial structure is of central importance in understanding the effects of fragmentation on population survival. Landscape spatial structure is the spatial relationships among habitat patches and the matrix in which they are embedded. Many general models of subdivided populations make the assumptions that (1) all habitat patches are equivalent in size and quality and (2) all local populations (in the patches) are equally accessible by dispersers. Models that gloss over spatial details of landscape structure can be useful for theoretical developments but will almost always be misleading when applied to real-world conservation problems. We show that local extinctions of fragmented populations are common. From this it follows that recolonization of local extinctions is critical for regional survival of fragmented populations. The probability of recolonization depends on (1) spatial relationships among landscape elements used by the population, including habitat patches for breeding and elements of the inter-patch matrix through which dispersers move, (2) dispersal characteristics of the organism of interest, and (3) temporal changes in the landscape structure. For endangered species, which are typically restricted in their dispersal range and in the kinds of habitat through which they can disperse, these factors are of primary importance and must be explicitly considered in management decisions.     

Patch Isolation, Corridor Effects, and Colonization by a Resident Sparrow in a Managed Pine Woodland

The isolation of habitat patches is often cited as having a major impact on the dynamics of small populations occupying patches in a complex landscape. Few studies, however, have provided field data demonstrating that isolation has an identifiable effect on specific populations independent of other factors such as local habitat quality or that landscape factors such as corridors can alleviate such effects. We conducted field surveys of Bachman's Sparrow ( Aimophila aestivalis ) populations in regions, which we call linear landscapes, where suitable habitat patches were isolated to varying degrees from potential sources of dispersing birds. In these linear landscapes, isolated patches of habitat were less likely to be colonized than were nonisolated patches. We also found that corridor configurations of habitat patches improved the ability of sparrows to find and settle in newly created patches. These results suggest that, for species that do not disperse easily through inhospitable landscapes, habitat occupancy at a regional scale can be enhanced by careful landscape design and planning.     

Importance of movement constraints in habitat selection studies

The aim of this study is to empirically illustrate the importance of taking movement constraints into account when testing for habitat selection with telemetry data. Global Positioning System relocations of two Scandinavian brown bears were used to compare the results of two different tests of habitat selection by the bears within their home range. Both relied on the comparison of observed dataset with datasets simulated under the hypothesis of random habitat use. The first analysis did not take movement constraints into account (simulations were carried out by randomly distributing a set of points in the home range) whereas the second analysis accounted for these constraints (simulations were carried out by building random trajectories in the home range). The results for the two analyses showed contrasted results. Therefore, not accounting for movement constraints in analyses may result in a misleading biological interpretation. Autocorrelation between relocations is not undesirable: it contains information about ecological processes that should be integrated in habitat selection analyses.     

Space use in lions, <Emphasis Type="Italic">Panthera leo</Emphasis>, in the Selous Game Reserve: social and ecological factors

Habitat qualities, such as food supply or access to refuges, often influence home-range size. Furthermore, such qualitative differences usually lead to conspecific competition over space, which can be an important factor in determining the distribution of individuals within populations. In carnivores, patterns of resource dispersion are hypothesized to determine home range-size and group size. But in contests over space (or other resources), larger groups usually dominate smaller ones, and group size should therefore also affect home-range size. Here I describe the space use of lions, Panthera leo, in the Selous Game Reserve, Tanzania, and ask whether space use is related to pride size, habitat, or relatedness. Home ranges varied in size, but size showed no correlation to number of adult females in the pride or to habitat type. Lions exhibited a significant preference for riverine and short-grass habitat, and a significant avoidance of acacia woodland. Habitat preference ratios largely reflected prey availability in each habitat. Outer areas, as well as core areas of home ranges, were often used by two or more prides. Overlaps showed no correlation to relatedness among prides or habitat type. Thus, whereas home-range sizes and overlaps were determined by factors that could not be revealed by demographic factors or analyses of habitat composition or genetic structure, lion space use within each home range seemed driven mostly by prey availability, which mainly varies with habitat type.     

Spatial memory, habitat auto-facilitation and the emergence of fractal home range patterns

Animals interact with their habitat in a manner which involves both negative and positive feedback mechanisms. We apply a specific modeling approach, “multi-scaled random walk”, for the scenario where a spatially explicit positive feedback process emerges from a combination of a spatial memory-dependent tendency to return to familiar patches and a consequently objective or subjective improvement of the quality of these patches (habitat auto-facilitation). In addition to the potential for local resource improvement from physically altering a patch, primarily known from the ecology of grazing ungulates, auto-facilitation from site fidelity may also embed more subtle subjective, individual-specific advantages from patch familiarity. Under the condition of resource superabundance, fitness gain from intra-home range patch fidelity creates a self-reinforcing use of the preferred patches on expense of a broader foraging in a priori equally favorable patches. Through this process, our simulations show that a spatially fractal dispersion of accumulated locations of the individual will emerge under the given model assumptions. Based on a conjecture that intra-home range patch fidelity depends on spatial memory we apply the multi-scaled random walk model to construct a spatially explicit habitat suitability parameter H_ij, which quantifies the dispersion of the generally most constraining resource from the individual's perspective. An intra-home range set of observed H-scores, H_obs, can then be estimated from a simple 2-scale calculation that is derived from the local dispersion of fixes. We show how the spatially explicit habitat utilization index H_obs not necessarily correlates positively with the local density fluctuations of fixes. The H-index solves some well-known problems from using the pattern of local densities of telemetry fixes - the classic utilization distribution - as a proxy variable for relative intra-home range habitat quality and resource selection. A pilot study on a set of telemetry fixes collected from a herd of free-ranging domestic sheep with overlapping summer home ranges illustrates how the H-index may be estimated and interpreted as a first-level approach towards a more extensive analysis of intra-home range habitat resource availability and patch preferences. Spatial memory in combination with site fidelity requires a modeling framework that explicitly describes the property of positive feedback mechanism under auto-facilitation in a spatio-temporally explicit manner.     

Costs and benefits of home range shifts among vervet monkeys (Cercopithecus aethiops) in Amboseli National Park,Kenya

Summary A decline in the density of trees and the deterioration of the habitat of vervet monkeys (Cercopithecus aethiops) in Amboseli National Park, Kenya, have been accompanied by movement of vervet groups into unfamiliar areas. These home range shifts were documented over a period of 26 months. The largest group moved into the home ranges of neighboring groups and acquired more trees, a resource that had become more limited. The groups whose home ranges were encroached upon did not share their home ranges with the intruders, but instead moved into areas that were unoccupied and had a lower density of trees. The largest group was thus more successful than smaller groups in competition for limited resources. For all but one group, movement into new areas was directly associated with the disappearances of female and immature monkeys. Most of the disappearances were attributable to predation. These results suggest that the cost of predation in unfamiliar areas may contribute to the evolution of philopatry in vervets. However, under unstable conditions, resource competition forces these animals to disperse (in groups) despite the high risk of predation. Offprint requests to: L.A. Isbell     

A Voronoi diagram based population model for social species of wildlife

A population model is presented that accounts for spatial structure within habitat patches. It is designed for social species of wildlife that form social group home ranges that are much smaller than patch size. The model represents social group home ranges by Voronoi regions that tessellate a patch to form a Voronoi diagram. Neighbouring social groups are linked with habitat-confined shortest paths and form a dispersal network. The model simulates population dynamics and makes use of Voronoi diagrams and dispersal networks as a spatial component. It then produces density maps as outputs. These are maps that show predicted animal densities across the patches of a landscape. A construction procedure for the particular Voronoi diagram type used by the model is described. As a test case, the model is run for the squirrel glider (Petaurus norfolcensis), a small arboreal marsupial native to Australia. A time series of density maps are produced that show squirrel glider density changing across a landscape through time.     

An Experimental Test of Matrix Permeability and Corridor Use by an Endemic Understory Bird

Abstract:  Because of widespread habitat fragmentation, maintenance of landscape connectivity has become a major focus of conservation planning, but empirical tests of animal movement in fragmented landscapes remain scarce. We conducted a translocation experiment to test the relative permeability of three landscape elements (open habitat, shrubby secondary vegetation, and wooded corridors) to movement by the Chucao Tapaculo ( Scelorchilus rubecula ), a forest understory bird endemic to South American temperate rainforest. Forty-one radio-tagged subjects were translocated (individually) to three landscape treatments consisting of small release patches that were either entirely surrounded by open habitat (pasture), entirely surrounded by dense shrubs, or linked to other patches by wooded corridors that were otherwise surrounded by open matrix. The number of days subjects remained in release patches before dispersal (a measure of habitat resistance) was significantly longer for patches surrounded by open habitat than for patches adjoining corridors or surrounded by dense shrubs. These results indicate that open habitat significantly constrains Chucao dispersal, in accord with expectation, but dispersal occurs equally well through wooded corridors and shrub-dominated matrix. Thus, corridor protection or restoration and management of vegetation in the matrix (to encourage animal movement) may be equally feasible alternatives for maintaining connectivity.     

Effectiveness of Corridors Relative to Enlargement of Habitat Patches

Abstract:  The establishment of biological corridors between two otherwise isolated habitat patches is a common yet contentious strategy for conserving populations in fragmented landscapes. We compared the effectiveness of corridors with the effectiveness of an alternate conservation strategy, the enlargement of existing habitat patches. We used a spatially explicit population model that simulated population size in two kinds of patches. One patch had a corridor that connected it to a larger "source" patch and the other patch was unconnected and enlarged at the periphery by an area the same size as the corridor. Patch isolation, corridor width, patch size, and the probability that individuals would cross the border from habitat to matrix were varied independently. In general, population size was greater in enlarged patches than in connected patches when patches were relatively large and isolated. Corridor width and the probability of crossing the border from habitat to matrix did not affect the relative benefit of corridors versus patch enlargement. Although biological corridors may mitigate potential effects of inbreeding depression at long time scales, our results suggest that they are not always the best method of conserving fragmented populations.     

Effects of temperature and precipitation on grassland bird nesting success as mediated by patch size

Grassland birds are declining faster than any other bird guild across North America. Shrinking ranges and population declines are attributed to widespread habitat loss and increasingly fragmented landscapes of agriculture and other land uses that are misaligned with grassland bird conservation. Concurrent with habitat loss and degradation, temperate grasslands have been disproportionally affected by climate change relative to most other terrestrial biomes. Distributions of grassland birds often correlate with gradients in climate, but few researchers have explored the consequences of weather on the demography of grassland birds inhabiting a range of grassland fragments. To do so, we modeled the effects of temperature and precipitation on nesting success rates of 12 grassland bird species inhabiting a range of grassland patches across North America (21,000 nests from 81 individual studies). Higher amounts of precipitation in the preceding year were associated with higher nesting success, but wetter conditions during the active breeding season reduced nesting success. Extremely cold or hot conditions during the early breeding season were associated with lower rates of nesting success. The direct and indirect influence of temperature and precipitation on nesting success was moderated by grassland patch size. The positive effects of precipitation in the preceding year on nesting success were strongest in relatively small grassland patches and had little effect in large patches. Conversely, warm temperatures reduced nesting success in small grassland patches but increased nesting success in large patches. Mechanisms underlying these differences may be patch‐size‐induced variation in microclimates and predator activity. Although the exact cause is unclear, large grassland patches, the most common metric of grassland conservation, appears to moderate the effects of weather on grassland‐bird demography and could be an effective component of climate‐change adaptation.