Using patch isolation metrics to predict animal movement in binary landscapes

Habitat isolation can affect the distribution and abundance of wildlife, but it is an ambiguous attribute to measure. Presumably, isolation is a characteristic of a habitat patch that reflects how spatially inaccessible it is to dispersing organisms. We identified four isolation metrics (nearest-neighbor distance, Voronoi polygons, proximity index, and habitat buffers) that were representative of the different families of metrics that are commonly used in the literature to measure patch isolation. Using simulated data, we evaluated the ability of each isolation metric to predict animal dispersal. We examined the simulated movement of organisms in two types of landscapes: an artificially-generated point-pattern landscapes where patch size and shape were consistent and only the arrangement of patches varied, and realistic landscapes derived from a geographic information system (GIS) of forest-vegetation maps where patch size, shape, and isolation were variable. We tested the performance of the four isolation metrics by examining the strength of the correlation between observed immigration rate in the simulations and each patch isolation metric. We also evaluated whether each isolation metric would perform consistently under varying conditions of patch size/shape, total amount of habitat in the landscape, and proximity of the patch to the landscape edge. The results indicate that a commonly-used distance-based metric, nearest-neighbor distance, did not adequately predict immigration rate when patch size and shape were variable. Area-informed isolation metrics, such as the amount of available habitat within a given radius of a patch, were most successful at predicting immigration. Overall, the use of area-informed metrics is advocated despite the limitation that these metrics require parameterization to reflect the movement capacity of the organism studied.This revised version was published online in May 2005 with corrections to the Cover Date.     

The relative impact of forest patch and landscape attributes on black howler monkey populations in the fragmented Lacandona rainforest, Mexico

Land-use change is forcing many animal populations to inhabit forest patches in which different processes can threaten their survival. Some threatening processes are mainly related to forest patch characteristics, but others depend principally on the landscape spatial context. Thus, the impact of both patch and landscape spatial attributes needs to be assessed to have a better understanding of the habitat spatial attributes that constraint the maintenance of populations in fragmented landscapes. Here, we evaluated the relative effect of three patch-scale (i.e., patch size, shape, and isolation) and five landscape-scale metrics (i.e., forest cover, fragmentation, edge density, mean inter-patch isolation distance, and matrix permeability) on population composition and structure of black howler monkeys (Alouatta pigra) in the Lacandona rainforest, Mexico. We measured the landscape-scale metrics at two spatial scales: within 100 and 500 ha landscapes. Our findings revealed that howler monkeys were more strongly affected by local-scale metrics. Smaller and more isolated forest patches showed a lower number of individuals but at higher densities. Population density also tended to be positively associated to matrices with higher proportion of secondary forests and arboreal crops (i.e. with greater permeability), most probably because these matrices can offer supplementary foods. The immature-to-female ratio also increased with matrix permeability, shape complexity, and edge density; habitat characteristics that can increase landscape connectivity and sources availability. The prevention of habitat loss and isolation, and the increment of matrix permeability are therefore needed for the conservation of this endangered Neotropical mammal.     

Isolation from forest reduces pollination,seed predation and insect scavenging in Swiss farmland

Habitat loss and fragmentation lead to changes in species richness and composition which may affect ecosystem services. Yet, few studies distinguish between the effects of habitat loss and isolation, or how multiple ecosystem services may be affected simultaneously. We investigated the effects of variation in cover of woody and open semi-natural habitats and isolation from forest on the relative functioning of pollination, seed predation and insect scavenging in agricultural landscapes. We established 30 sites in grassland locations in the Swiss plateau around Berne. The sites varied independently in their isolation from forest edges, in the percentage of woody habitats and in the percentage of open semi-natural habitats in the surrounding landscape (500 m radius). We experimentally exposed primroses, sunflower seeds and cricket corpses during spring 2008. None of the three studied services was affected by variation in woody or open semi-natural habitat cover. However, the proportion of flowers setting seed was significantly reduced by isolation from forest. Further, seed predation and insect scavenging were significantly lower at isolated sites than at sites connected to woody habitat. This pattern was particularly pronounced for seeds and insect corpses that were enclosed by wire netting and thus inaccessible to vertebrates. Thus, all three studied services responded quite similarly to the landscape context. The observed small-scale determination of seed set, seed predation and insect scavenging contrasts with larger-scale determination of pollination and insect pest control found in other studies.     

Spatial configuration matters: a test of the habitat amount hypothesis for plants in calcareous grasslands

Context

A recent hypothesis, the habitat amount hypothesis, predicts that the total amount of habitat in the landscape can replace habitat patch size and isolation in studies of species richness in fragmented landscapes.

Objectives

To test the habitat amount hypothesis by first evaluating at which spatial scale the relationship between species richness in equal-sized sample quadrats and habitat amount was the strongest, and then test the importance of spatial configuration of habitat—measured as local patch size and isolation—when habitat amount was taken into account.

Methods

A quasi-experimental setup with 20 habitat patches of dry calcareous grasslands varying in patch size, patch isolation and habitat amount at the landscape scale was established in the inner Oslo fjord, Southern Norway. We recorded species richness of habitat specialists of vascular plants in equal-sized sample quadrats and analysed the relationship between species richness, habitat amount in the landscape and patch size and isolation.

Results

Although the total amount of habitat in a 3 km-radius around the local patch was positively related to species richness in the sample quadrats, local patch size had an additional positive effect, and the effect of patch size was higher when the amount of habitat within the 3 km-radius was high than when it was low.

Conclusions

In our study system of specialist vascular plants in dry calcareous grasslands, we do not find support for the habitat amount hypothesis.

     

Watershed analysis of forest fragmentation by clearcuts and roads in a Wyoming forest

Remotely sensed data and a Geographic Information System were used to compare the effects of clearcutting and road-building on the landscape pattern of the Bighorn National Forest, in north-central Wyoming. Landscape patterns were quantified for each of 12 watersheds on a series of four maps that differed only in the degree of clearcutting and road density. We analyzed several landscape pattern metrics for the landscape as a whole and for the lodgepole pine and spruce/fir cover classes across these maps, and determined the relative effects of clearcutting and road building on the pattern of each watershed. At both the landscape- and cover class-scales, clearcutting and road building resulted in increased fragmentation as represented by a distinct suite of landscape structural changes. Patch core area and mean patch size decreased, and edge density and patch density increased as a result of clearcuts and roads. Clearcuts and roads simplified patch shapes at the landscape scale, but increased the complexity of lodgepole pine patches. Roads appeared to be a more significant agent of change than clearcuts, and roads which were more evenly distributed across a watershed had a greater effect on landscape pattern than did those which were densely clustered. Examining individual watersheds allows for the comparison of fragmentation among watersheds, as well as across the landscape as a whole. Similar studies of landscape structure in other National Forests and on other public lands may help to identify and prevent further fragmentation of these areas.     

Soil nitrogen and carbon dynamics in a fragmented landscape experiencing forest succession

Forest fragmentation is an increasingly common feature across the globe, but few studies examine its influence on biogeochemical fluxes. We assessed the influence of differences in successional trajectory and stem density with forest patch size on biomass quantity and quality and N transformations in the soil at an experimentally fragmented landscape in Kansas, USA. We measured N-related fluxes in the laboratory, not the field, to separate effects of microclimate and fragment edges from the effects of inherent biomass differences with patch size. We measured net N mineralization and N₂O fluxes in soil incubations, gross rates of ammonification and nitrification, and microbial biomass in soils. We also measured root and litterfall biomass, C:N ratios, and δ¹³C and δ¹⁵N signatures; litterfall [cellulose] and [lignin]; and [C], [N], and δ¹³C and δ¹⁵N of soil organic matter. Rates of net N mineralization and N₂O fluxes were greater (by 113% and 156%, respectively) in small patches than in large, as were gross rates of nitrification. These differences were associated with greater quantities of root biomass in small patch soil profiles (664.2 ± 233.3 vs 192.4 ± 66.2 g m⁻² for the top 15 cm). These roots had greater N concentration than in large patches, likely generating greater root derived organic N pools in small patches. These data suggest greater rates of N cycling in small forested patches compared to large patches, and that gaseous N loss from the ecosystem may be related to forest patch size. The study indicates that the differences in successional trajectory with forest patch size can impart significant influence on soil N transformations in fragmented, aggrading woodlands.     

Statistical models of landscape pattern metrics,with applications to regional scale dynamic forest simulations

Forest managers in Canada need to model landscape pattern or spatial configurationoverlarge (100,000 km²) regions. This presents a scalingproblem, as landscape configuration is measured at a high spatial resolution,but a low spatial resolution is indicated for regional simulation. We present astatistical solution to this scaling problem by showing how a wide range oflandscape pattern metrics can be modelled from low resolution data. Our studyarea comprises about 75,000 km² of boreal mixedwoodforest in northeast Alberta, Canada. Within this area we gridded a sample of 84digital forest cover maps, each about 9500 ha in size, to aresolution of 1 ha and used FRAGSTATS to compute a suite oflandscape pattern metrics for each map. We then used multivariate dimensionreduction techniques and canonical correlation analysis to model therelationship between landscape pattern metrics and simpler stand table metricsthat are easily obtained from non-spatial forest inventories. These analyseswere performed on four habitat types common in boreal mixedwood forests: youngdeciduous, old deciduous, white spruce, and mixedwood types. Using only threelandscape variables obtained directly from stand attribute tables (totalhabitatarea, and the mean and standard deviation of habitat patch size), ourstatistical models explained more than 73% of the joint variation in fivelandscape pattern metrics (representing patch shape, forest interior habitat,and patch isolation). By PCA, these five indices captured much of the totalvariability in the rich set of landscape pattern metrics that FRAGSTATS cangenerate. The predictor variables and strengths of association were highlyconsistent across habitat classes. We illustrate the potential use of suchstatistical relationships by simulating the regional, cumulative effects ofwildfire and forest management on the spatial arrangement of forest patches,using non-spatial stand attribute tables.This revised version was published online in May 2005 with corrections to the Cover Date.     

An early synthesis of the habitat amount hypothesis

Context

The ecological literature is filled with studies highlighting the importance of both habitat loss and fragmentation on biodiversity. The patch concept has been central to these findings, being also at the heart of many ecological theories. Recently, the habitat amount hypothesis has been proposed as an alternative, where the patch concept is put to a rest, and both patch size and patch isolation effects on species richness are reduced to a single gradient: habitat loss in the landscape.

Objectives

As this theory stated clear predictions that could be experimentally tested, many formal tests of the hypothesis have been published recently and this study aims at synthesizing their results.

Methods

A meta-analysis of 13 tests of the habitat amount hypothesis was conducted, to produce a single combined test of the theory.

Results

The 13 tests combined suggest that effects of patch size and isolation, while controlling for habitat amounts, do exist although their overall effect is weak (r?=?0.158).

Conclusions

Literal interpretations of the habitat amount hypothesis, where patch size and isolation have absolutely no effect on species richness, are probably oversimplifications of the processes at work. Still, the theory could prove useful as a baseline of the effects of habitat loss, against which patch size and isolation effects must be contrasted.

     

Landscape Fragmentation and Ice Storm Damage in Eastern Ontario Forests

With return times between 20 and 100 years, ice storms are a primary disturbance type for temperate forests of eastern North America. Many studies have been conducted at the forest patch and plot scales to examine relations between damage and variables describing site, composition and structure. This paper presents results from a landscape scale study of fragmentation relations with damage in eastern Ontario forests. Data previously collected for two independent and spatially non-overlapping patch level damage studies were used. A Generalized Linear Model (GLM) was used to analyse relations between damage and fragmentation metrics representing patch isolation, edge density, and the relative size and distribution of patches in the landscape. The metrics were applied using spatial extents of 1 × 1 km and 4 × 4 km, following analyses of the variability of numbers of patches and of the lacunarity of forest patterns over a range of extents. The results showed that patch isolation, as measured by the mean Euclidean distance between patches (ENN) was significantly related to damage.     

Evaluation of patch isolation metrics in mosaic landscapes for specialist vs. generalist dispersers

We examined the effects of matrix structure and movement responses of organisms on the relationships between 7 patch isolation metrics and patch immigration. Our analysis was based on simulating movement behaviour of two generic disperser types (specialist and generalist) across mosaic landscapes containing three landcover types: habitat, hospitable matrix and inhospitable matrix. Movement, mortality and boundary crossing probabilities of simulated individuals were linked to the landcover and boundary types in the landscapes. The results indicated that area-based isolation metrics generally predict patch immigration more reliably than distance-based isolation metrics. Relationships between patch isolation metrics and patch immigration varied between the two generic disperser types and were affected by matrix composition or matrix fragmentation. Patch immigration was always affected by matrix composition but not by matrix fragmentation. Our results do not encourage the generic use of patch isolation metrics as a substitute for patch immigration, in particular in metapopulation models where generic use may result in wrong projections of the survival probability of metapopulations. However, our examination of the factors affecting the predictive potential of patch isolation metrics should facilitate interpretation and comparison of existing patch isolation studies. Future patch isolation studies should include information about landscape structure and the dispersal distance and dispersal behaviour of the organism of interest.This revised version was published online in May 2005 with corrections to the Cover Date.     

Transitions in forest fragmentation: implications for restoration opportunities at regional scales

Where the potential natural vegetation is continuous forest (e.g., eastern US), a region can be divided into smaller units (e.g., counties, watersheds), and a graph of the proportion of forest in the largest patch versus the proportion in anthropogenic cover can be used as an index of forest fragmentation. If forests are not fragmented beyond that converted to anthropogenic cover, there would be only one patch in the unit and its proportional size would equal 1 minus the percentage of anthropogenic cover. For a set of 130 watersheds in the mid-Atlantic region, there was a transition in forest fragmentation between 15 and 20% anthropogenic cover. The potential for mitigating fragmentation by connecting two or more disjunct forest patches was low when percent anthropogenic cover was low, highest at moderate proportions of anthropogenic cover, and again low as the proportion of anthropogenic cover increased toward 100%. This fragmentation index could be used to prioritize locations for restoration by targeting watersheds where there would be the greatest increase in the size of the largest forest patch.     

Matrix is important for mammals in landscapes with small amounts of native forest habitat

Acknowledgment that the matrix matters in conserving wildlife in human-modified landscapes is increasing. However, the complex interactions of habitat loss, habitat fragmentation, habitat condition and land use have confounded attempts to disentangle the relative importance of properties of the landscape mosaic, including the matrix. To this end, we controlled for the amount of remnant forest habitat and the level of fragmentation to examine mammal species richness in human-modified landscapes of varying levels of matrix development intensity and patch attributes. We postulated seven alternative models of various patch habitat, landscape and matrix influences on mammal species richness and then tested these models using generalized linear mixed-effects models within an information theoretic framework. Matrix attributes were the most important determinants of terrestrial mammal species richness; matrix development intensity had a strong negative effect and vegetation structural complexity of the matrix had a strong positive effect. Distance to the nearest remnant forest habitat was relatively unimportant. Matrix habitat attributes are potentially a more important indicator of isolation of remnant forest patches than measures of distance to the nearest patch. We conclude that a structurally complex matrix within a human-modified landscape can provide supplementary habitat resources and increase the probability of movement across the landscape, thereby increasing mammal species richness in modified landscapes.     

On the seasonal effect of landscape structure on a bird species: the thorn-tailed rayadito in a relict forest in northern Chile

Forest bird species exhibit noticeable seasonal behavioral changes that might lead to contrasting effects of landscape pattern upon species abundance and performance. We assessed if the effect of patch and habitat attributes on the landscape use of thorn-tailed rayaditos (Aphrastura spinicauda), a forest bird in a relict patchy forest in northern Chile, varied temporally in association with changes in the behavior of individuals linked to breeding vs. non-breeding conditions. We also assessed the relationship between nest success and patch and habitat attributes, as nest success might be associated to the density rayaditos during the breeding season. We found that density of rayaditos was affected by patch size and functional connectivity but not by habitat structure and that the magnitude of the effect of patch size was greater during the non-breeding season, thus supporting the existence of a temporally variable effect of landscape pattern. Similarly, the nest success of rayaditos was positively affected by functional connectivity and negatively by structural connectivity. We hypothesize that these results emerged from the interaction among territorial behavior, resource limitation and predation risk. Despite the variable intensity of the effect of patch size upon density, however, this landscape attribute, in addition to connectivity, is essential for the persistence of rayaditos at this relict patchy forest landscapes.     

The importance of integrating landscape ecology in habitat models: isolation-driven occurrence of north island robins in a fragmented landscape

Although the role of habitat fragmentation in species declines is well recognised, the effect of habitat quality on species distributions is often studied using presence–absence models that ignore metapopulation dynamics. We compared three approaches to model the presence–absence of North Island robins in 400 sites among 74 fragments of native forest in a 15,000-ha agricultural landscape in New Zealand. The first approach only considered local habitat characteristics, the second approach only considered metapopulation factors (patch size and isolation), and the third approach combined these two types of factors. The distribution of North Island robins was best predicted by patch isolation, as their probability of occurrence was negatively correlated with isolation from neighbouring patches and from the closest major forests, which probably acted as a source of immigrants. The inclusion of habitat factors gave only a slight increase in predictive power and indicated that robins were more likely to occur in areas with tall canopy, tall understory and low density of young trees. We modelled the effect of isolation using an index of functional patch connectivity based on dispersal behaviour of radio-tracked juveniles, and this functional index greatly improved the models in comparison to classical indices relying on Euclidean distances. This study highlights the need to incorporate functional indices of isolation in presence–absence models in fragmented landscapes, as species occurrence can otherwise be a misleading predictor of habitat quality and lead to wrong interpretations and management recommendations.     

Relative importance of habitat area and isolation for bird occurrence patterns in a naturally patchy landscape

There is debate among ecologists about whether total habitat area or patch arrangement contributes most to population and/or community responses to fragmented or patchy landscapes. We tested the relative effects of patch area and isolation for predicting bird occurrence in a naturally patchy landscape in the Bear River Mountains of Northern Utah, USA. We selected focal patches (mountain meadows) ranging in elevation from 1,920 to 2,860 m and in size from 0.6 to 182 ha. Breeding birds were sampled in each focal meadow during the summers of 2003 and 2004 using variable-distance point transects. Logistic regression and likelihood-based model selection were used to determine the relationship between likelihood of occurrence of three bird species (Brewer’s sparrow, vesper sparrow, and white-crowned sparrow) and area, isolation, and proximity metrics. We used model weights and model-averaged confidence intervals to assess the importance of each predictor variable. Plots of area versus isolation were used to evaluate complex relationships between the variables. We found that meadow area was the most important variable for explaining occurrence for two species, and that isolation was the most important for the other. We also found that the absolute distance was more appropriate for evaluating isolation responses than was the species-specific proximity metric. Our findings add clarity to the debate between ecologists regarding the relative importance of area and isolation in species responses to patchy landscapes.     

Creating landscape patterns by forest cutting: Ecological consequences and principles

Landscape structural characteristics, such as patch size, edge length, and configuration, are altered markedly when management regimes are imposed on primeval landscapes. The ecological consequences of clearcutting patterns were explored by using a model of the dispersed patch or checkerboard system currently practiced on federal forest lands in the western United States. Thresholds in landscape structure were observed on a gradient of percentages of landscape cutover. Probability of disturbance,e.g., wildfire and windthrow, and biotic components,e.g., species diversity and game populations, are highly sensitive to these structural changes. Altering the spatial configuration and size of clearcuts provides an opportunity to create alternative landscapes that differ significantly in their ecological characteristics. Both ecosystem and heterogeneous landscape perspectives are critical in resource management.     

Does matrix resistance influence Red squirrel (Sciurus vulgaris L. 1758) distribution in an urban landscape?

In determining isolation effects in fragmented populations, the landscape matrix is not often considered. Usually simple distance measures are used to quantify degree of isolation. We tested the effect of the matrix on the presence of red squirrels in 354 wooded patches in the Brussels Region, by comparing several isolation measures. These were 1) distance to the nearest source patch, 2) the Hanski-measure (a combination of distance to and size of all possible sources), 3) effective distances calculated from different least cost models using the ArcView grid extension ‘Cost Distance’ (a combination of distance and resistance of the landscape, with different resistances for different landscape types) and 4) some combinations of the Hanski-measure and the effective distances. Size and quality of the target patches were always included in the tests of the predictive power of different isolation measures on squirrel presence/absence. All variables examined (patch size, quality and isolation) significantly influenced squirrel presence. Models using the effective distances gave the best results. Models including the Hanski-measure improved significantly when Euclidean distance was replaced by effective distance, showing that parameterisation of matrix resistance added significant additional explanatory power when modelling squirrel presence. This revised version was published online in July 2006 with corrections to the Cover Date.     

城市森林景观动态变化与驱动力分析——以沈阳市铁西区为例

在遥感和GIS技术的支持下,借助各种遥感影像,采用转移矩阵和景观斑块变化类型的分析方法,对2001～2011年沈阳市铁西区大中型(面积大于400m2)城市森林景观动态变化进行分析。结果表明:10年间,铁西区各类型用地相互转化频繁,大中型城市森林斑块总面积和总数量都呈现先增后减的趋势,斑块整体水平向中小型化发展。而工业用地、企事业单位用地和其他用地中的斑块更新度高。研究区域以工业迁移因素为核心,经济发展、人口增长及管理因素成为景观动态变化的主要驱动力。     

Comparing the landscape level perceptual abilities of forest sciurids in fragmented agricultural landscapes*

Perceptual range is the maximum distance from which an animal can perceive the presence of remote landscape elements such as patches of habitat. Such perceptual abilities are of interest because they influence the probability that an animal will successfully disperse to a new patch in a landscape. Furthermore, understanding how perceptual range differs between species may help to explain differential species sensitivity to patch isolation. The objective of this research was to assess the perceptual range of eastern chipmunks (Tamias striatus), gray squirrels (Sciurus carolinensis), and fox squirrels (Sciurus niger) in fragmented agricultural landscapes. Animals were captured in remote woodlots and translocated to unfamiliar agricultural fields. There they were released at different distances from a woodlot and their movements towards or away from the woodlot were used to assess their ability to perceive forested habitat. Observed perceptual ranges of approximately 120 m for chipmunks, 300 m for gray squirrels, and 400 m for fox squirrels, suggest that differences in landscape-level perceptual abilities may influence the occurrence of these species in isolated habitat patches.     

Designing for conservation outcomes: the value of remnant habitat for reptiles on ski runs in subalpine landscapes

Subalpine ecosystems are centres of endemism that are important for biodiversity. However, these areas are under threat from the creation, expansion and continued modification of ski runs, activities that have largely negative effects on wildlife. Despite this threat, research on the impacts of ski runs is limited for reptiles—particularly regarding the value of remnant vegetation retained on ski runs. Here we quantify the effects of habitat loss and fragmentation (i.e., patch size, patch isolation and edge effects) on the abundance of a common subalpine lizard and on thermal regimes (a key determinant of lizard distribution) in an Australian ski resort. The number of lizards observed differed significantly with habitat type (ski runs vs. forested areas) and patch isolation, but not patch size. In addition, the edges of patches supported more lizards than any other habitat type. These patterns of lizard distribution can be explained, in part, by the differing thermal regimes in each habitat. Ski runs had significantly higher ground surface temperatures than any other habitat type, precluding their use for a considerable proportion of the activity period of a lizard. In comparison, edges were characterised by lower temperatures than ski runs, but higher temperatures than the core of forested areas, potentially providing a favourable environment for thermoregulation. Based on our results, we conclude that although modified ski runs have a negative effect on lizards, patches of remnant vegetation retained on ski runs are of value for reptiles and their conservation could help mitigate the negative effects of habitat loss caused by ski run creation.