The behavior of landscape metrics commonly used in the study of habitat fragmentation

A meaningful interpretation of landscape metrics is possible only when the limitations of each measure are fully understood, the range of attainable values is known, and the user is aware of potential shifts in the range of values due to characteristics of landscape patches. To examine the behavior of landscape metrics, we generated artificial landscapes that mimicked fragmentation processes while controlling the size and shape of patches in the landscape and the mode of disturbance growth. We developed nine series of increasingly fragmented landscapes and used these to investigate the behavior of edge density, contagion, mean nearest neighbor distance, mean proximity index, perimeter-area fractal dimension, and mass fractal dimension. We found that most of the measures were highly correlated, especially contagion and edge density, which had a near-perfect inverse correspondence. Many of the measures were linearly-associated with increasing disturbance until the proportion of disturbance on the landscape was approximately 0.40, with non-linear associations at higher proportions. None of the measures was able to differentiate between landscape patterns characterized by dispersed versus aggregated patches. The highest attainable value of each measure was altered by either patch size or shape, and in some cases, by both attributes. We summarize our findings by discussing the utility of each metric.     

Effects of habitat fragmentation on four rodent species in a Polish farm landscape

Bank vole, striped field mouse, wood mouse, and yellow-necked mouse populations were studied in a mosaic of field and forest habitats. Live-trapping was carried out in 8 woodlots of different sizes (1.5–9.5 ha), situated 5 to 900 m from each other and surrounded by agricultural fields. Near the study area a dense, several hundred hectare forest complex was situated. It was found that the densities of all the studied species' populations in the woodlots were positively correlated with woodlot quality. For local bank vole populations a positive correlation of density with the surface area and circumference of woodlots, as well as with the area/circumference ratio was found. A negative correlation was found for population density and the distance between a given woodlot and the forest complex. For the yellow-necked mouse a positive correlation occurred between the density of local populations and the distance to the nearest neighboring woodlot. For the striped field mouse a positive correlation was found only between the population density in each woodlot and the distance to the forest complex. The wood mouse was insensitive to the variations in woodlot features present except for woodlot quality, and hence was probably responding in density to some other factors. Four rodent species, coexisting in the field-forest habitat mosaic demonstrated different reactions to its spatial characteristics, which were mainly related to different habitat preferences, spatial behavior, and mobility of individuals of the studied species.     

Effects of habitat fragmentation on Eurasian red squirrel (Sciurus vulgaris) in a forest landscape

We studied the effects of habitat fragmentation, measured as forest stand size and isolation, on the distribution of Eurasian red squirrels (Sciurus vulgaris). Squirrel density was surveyed during four years in 46 forest stands (0.1–500 ha) in a forest landscape in south-central Sweden. The only factor that significantly influenced a density index was the proportion of spruce within a habitat fragment. Neither fragment size nor degree of isolation were significant. Furthermore, none of the interactions with year were significant, suggesting the same pattern in all four years. Thus, the effect of habitat fragmentation in this study seems to be only pure habitat loss, i.e. halving the proportion of preferred habitat in the landscape should result in a halving of the red squirrel population. Therefore, the landscape can be viewed as functionally continuous for the squirrels, although the preferred habitat was divided into fragments. The most likely explanation for the difference between this study and other studies on squirrels that found effects due to habitat fragmentation is a combination of shorter distances and less hostile surroundings in our study area. To identify landscape effects requires multiple studies because single studies usually consider only one landscape.     

Separating the effects of habitat area,fragmentation and matrix resistance on genetic differentiation in complex landscapes

Little is known about how variation in landscape mosaics affects genetic differentiation. The goal of this paper is to quantify the relative importance of habitat area and configuration, as well as the contrast in resistance between habitat and non-habitat, on genetic differentiation. We hypothesized that habitat configuration would be more influential than habitat area in influencing genetic differentiation. Population size is positively related to habitat area, and therefore habitat area should affect genetic drift, but not gene flow. In contrast, differential rates and patterns of gene flow across a landscape should be related to habitat configuration. Using spatially explicit, individual-based simulation modeling, we found that habitat configuration had stronger relationships with genetic differentiation than did habitat area, but there was a high degree of confounding between the effects of habitat area and configuration. We evaluated the predictive ability of six widely used landscape metrics and found that patch cohesion and correlation length of habitat are among the strongest individual predictors of genetic differentiation. Correlation length, patch density and clumpy are the most parsimonious set of variables to predict the magnitude of genetic differentiation in complex landscapes.     

Effects of habitat fragmentation on meadow vole (Microtus pennsylvanicus) population dynamics in experiment landscape patches

This study examined the effects of habitat fragmentation on meadow vole (Microtus pennsylvanicus) population dynamics in experimental landscape patches. The study was conducted from May–November 1993 at the Miami University Ecology Research Center. Eight 0.1-ha small mammal enclosures were used. Four enclosures contained a 160 m² nonfragmented patch and four enclosures contained four 40 m² fragmented patches. Thus, each treatment was replicated 4 times in a systematic research design. The patches in both treatments contained high-quality habitat surrounded by low-quality matrix. Six pairs of adult meadow voles were released in each enclosure on 27 May 1993. Populations were monitored by live-trapping and radio-telemetry methods. Significantly greater densities of female voles were found during October in the fragmented treatment compared to the nonfragmented treatment. Also, significantly more females than males were found in the fragmented treatment compared to the nonfragmented treatment for the total study period. Significantly more subadult and juvenile males were found in the matrix versus the patch of the nonfragmented treatment compared to the fragmented treatment. Males in the fragmented treatment had significantly greater mean home range size than males or females in the nonfragmented treatment. There appears to exist a relationship between patch fragmentation and the social structure of meadow vole populations; this relationship appears to function as a population regulatory mechanism.     

Functional rather than structural connectivity explains grassland plant diversity patterns following landscape scale habitat loss

Context

Functional connectivity is vital for plant species dispersal, but little is known about how habitat loss and the presence of green infrastructure interact to affect both functional and structural connectivity, and the impacts of each on species groups.

Objectives

We investigate how changes in the spatial configuration of species-rich grasslands and related green infrastructure such as road verges, hedgerows and forest borders in three European countries have influenced landscape connectivity, and the effects on grassland plant biodiversity.

Methods

We mapped past and present land use for 36 landscapes in Belgium, Germany and Sweden, to estimate connectivity based on simple habitat spatial configuration (structural connectivity) and accounting for effective dispersal and establishment (functional connectivity) around focal grasslands. We used the resulting measures of landscape change to interpret patterns in plant communities.

Results

Increased presence of landscape connecting elements could not compensate for large scale losses of grassland area resulting in substantial declines in structural and functional connectivity. Generalist species were negatively affected by connectivity, and responded most strongly to structural connectivity, while functional connectivity determined the occurrence of grassland specialists in focal grasslands. Restored patches had more generalist species, and a lower density of grassland specialist species than ancient patches.

Conclusions

Protecting both species rich grasslands and dispersal pathways within landscapes is essential for maintaining grassland biodiversity. Our results show that increases in green infrastructure have not been sufficient to offset loss of semi-natural habitat, and that landscape links must be functionally effective in order to contribute to grassland diversity.

     

A controlled,hierarchical study of habitat fragmentation: responses at the individual,patch, and landscape scale

We compared the performance of individuals and whole populations of meadow voles, Microtus pennsylvanicus, within and between experimentally created habitat fragments of three sizes (1.0, 0.25, and 0.0625 ha) and between a 20-ha fragmented and a 20-ha continuous habitat landscape. We recorded 10,020 captures of 3946 individuals over 17 censuses between June 1993 and October 1994. Five demographic parameters showed significantly different population responses between the two landscapes but no difference in tests comparing fragment size: i.e., mean and peak population densities (the latter, in each of the two growing seasons) averaged 149 to 172% higher, population growth rate averaged 219% higher, and adult recruitment 170% higher in fragmented than in the continuous control landscape. Observations at the individual level (body sizes, rates of reproduction, residence times) suggested that these landscape differences involved enhanced performance of adult females associated with edge habitats rather than differential immigration or emigration. If this turns out to be a common response to fragmentation, the detection of such responses will be greater when comparing fragmented and unfragmented landscapes with qualitatively different structure than for fragments of varied size with differing proportions of edge. That responses to habitat fragmentation may be more evident at the very small (individual) and very large (landscape) scales, but may be obscured at the intermediate spatial scale of fragments, is a proposition that clearly requires more attention.     

Testing landscape metrics as indicators of habitat loss and fragmentation in continuous eucalypt forests (Queensland,Australia)

Landscape metrics are widely applied in landscape ecology to quantify landscape structure. However, many are poorly tested and require rigorous validation if they are to serve as reliable indicators of habitat loss and frag-mentation, such as Montreal Process Indicator 1.1e. We apply a landscape ecology theory, supported by explor-atory and confirmatory statistical techniques, to empirically test landscape metrics for reporting Montreal Process Indicator 1.1e in continuous dry eucalypt forests of sub-tropical Queensland, Australia. Target biota examined included: the Yellow-bellied Glider (Petaurus australis); the diversity of nectar and sap feeding glider species including P. australis, the Sugar Glider P. breviceps, the Squirrel Glider P. norfolcensis, and the Feathertail GliderAcrobates pygmaeus; six diurnal forest birds species; total diurnal bird species diversity; and the density of nec-tar-feeding diurnal bird species. Two scales of influence were considered: the stand-scale (2 ha), and a series of radial landscape extents (500 m –2 km;78–1250 ha) surrounding each fauna transect. For all biota, stand-scale structural and compositional attributes were found to be more influential than landscape metrics. For the Yellow-belliedGlider, the proportion of trace habitats with a residual element of old spotted-gum/ironbark eucalypt trees was a significant landscape metric at the 2 km landscape extent. This is a measure of habitat loss rather than habitat fragmentation. For the diversity of nectar and sap feeding glider species, the proportion of trace habitats with a high coefficient of variation in patch size at the 750 m extent was a significant landscape metric. None of the landscape metrics tested was important for diurnal forest birds. We conclude that no single landscape metricadequately captures the response of the regions forest biota per se. This poses a major challenge to regional reporting of Montreal Process Indicator 1.1e, fragmentation of forest types.This revised version was published online in May 2005 with corrections to the Cover Date.     

Surrounding landscape structures,rather than habitat age,drive genetic variation of typical calcareous grassland plant species

Landscape Ecology - Land use change reduced calcareous grasslands throughout Europe during the last decades. Subsequent fragmentation and habitat deterioration led, moreover, to a massive...     

The effects of future urban development on habitat fragmentation in the Santa Monica Mountains

A site suitability model of urban development was created for the Santa Monica Mountains in southern California, USA, to project to what degree future development might fragment the natural habitat. The purpose was to help prioritize land acquisition for the Santa Monica Mountains National Recreation Area and examine to what extent projected urban development would affect distinct vegetation classes. The model included both environmental constraints (slope angle), and spatial factors related to urban planning (proximity to roads and existing development, proposed development, and areas zoned for development). It implemented a stochastic component; areas projected to have high development potential in the suitability model were randomly selected for development. Ownership tracts were used as the spatial unit of development in order to give the model spatial realism and not arbitrarily `develop' grid cells. Using different assumptions and parameters, the model projected the pattern of development from 5 to 25 years hence (based on recent development rates in the area). While <25% of the remaining natural landscape is removed under these scenarios, up to 30% of core (interior) habitat area is lost and edge length between natural vegetation and development increases as much as 45%. Measures of landscape shape complexity increased with area developed and number of patches of natural habitat increased four- to nine-fold, depending upon model parameters. This increase in fragmentation occurs because of the existing patterns of land ownership, where private (`developable') land is interspersed with preserved park lands.     

Evidence for a direct negative effect of habitat fragmentation on forest herb functional diversity

The effects of habitat fragmentation on species richness and composition have been extensively studied. However, little is known about how fragmentation affects functional diversity patterns. Fragmentation can indeed affect functional diversity directly (e.g. by promoting traits associated to long-distance dispersal when fragment isolation increases) or indirectly (e.g. by decreasing species richness, hence trait diversity, when fragment area decreases). Here, we used structural equation modeling to determine whether factors associated to forest fragmentation, namely area, habitat heterogeneity, spatial isolation and age have a direct effect on forest herb functional diversity. Using occurrence data from 243 forest fragments located in northern France and six plant life-history traits, we estimated species richness and calculated functional diversity in each of these 243 forest fragments. We found that species richness was the primary driver of functional diversity in these fragments, with a strong positive and direct relationship between species richness and functional diversity. Interestingly, both fragment isolation and age had a direct negative effect on functional diversity independent of their effects on species richness. Isolation selected life-history traits associated with long-distance dispersal, while age selected for life-history traits typical of forest habitat specialists. Isolated and/or older forest fragments are thus at greater risk of local species and functional extinctions, and hence making these forest fragments particularly vulnerable to future global changes.     

Estimating the consequences of habitat fragmentation on extinction risk in dynamic landscapes

Analyzing the population dynamic consequences of spatio-temporal changes in landscape structure is a formidable challenge for spatial ecology. One key population dynamic process in fragmented landscapes is the influence of isolation on colonization rate and thereby on the occurrence of species in habitat fragments, but it is not obvious how isolation should be measured in landscapes that are affected by on-going habitat loss and fragmentation. We suggest the following procedure for the measurement of spatio-temporal isolation. First, a historical record of habitat loss and fragmentation in the landscape is prepared based on snapshots of the extent of the suitable habitat for the focal species. Second, a metapopulation model is used to simulate the occurrence of the species in this landscape, assuming the empirically observed landscape change. The model-predicted pattern of habitat occupancy at a particular point in time (usually the present time) is then compared with empirical observations on the occurrence of the species. We describe a metapopulation model that has been constructed for this purpose, and we apply it to a changing landscape of boreal forests in eastern Finland. We give an example on the occurrence of four threatened polyporous fungi in 18 small fragments of old-growth forest. In none of the species does the current isolation of the fragments nor the time since their isolation explain the occurrence of the species in the study fragments, but in three species the model-predicted occupancy probability had a significant effect on the observed abundance of the species. The model-predicted occupancy probabilities were also calculated by ignoring past landscape changes, that is, by assuming that the landscape had remained in the present configuration for a long time. These probabilities had a significant effect on the abundance of only one of the four species, suggesting that the occurrence of the species tracks landscapes changes with a noticable time lag.This revised version was published online in May 2005 with corrections to the Cover Date.     

The impact of habitat fragmentation on the ecology of xenarthrans (Mammalia) in the Brazilian Cerrado

Landscape Ecology - The impact of deforestation and fragmentation upon ecologically important and poorly known groups is currently an important issue for conservation biology. Herein we describe...     

An experimental test of the habitat amount hypothesis reveals little effect of habitat area but transient or indirect effects of fragmentation on local species richness

Landscape Ecology - The habitat amount hypothesis (HAH) posits that local species richness is driven more by the amount of habitat in the surrounding landscape than by local patch size or habitat...     

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.     

The risk of ignoring fear: underestimating the effects of habitat loss and fragmentation on biodiversity

Landscape Ecology - Habitat loss and fragmentation threaten species not only through structural landscape changes and resource reduction, but also through modifications to species’...     

Building patterns and landscape fragmentation in northern Wisconsin,USA

Housing growth is prevalent in rural areas in the United States and landscape fragmentation is one of its many effects. Since the 1930s, rural sprawl has been increasing in areas rich in recreational amenities. The question is how housing growth has affected landscape fragmentation. We thus tested three hypotheses relating land cover and land ownership to density and spatial pattern of buildings, and examined whether building density or spatial pattern of buildings was a better predictor for landscape fragmentation. Housing locations were mapped from 117 1:24,000-scale USGS topographic maps across northern Wisconsin. Patch-level landscape metrics were calculated on the terrestrial area remaining after applying 50, 100 and 250 m disturbance zones around each building. Our results showed that building density and the spatial pattern of buildings were affected mostly by lake area, public land ownership, and the abundance of coniferous forest, agricultural land, and grassland. A full 40% of the houses were within 100 m of lakeshores. The clustering of buildings within 100 m of lakeshores limited fragmentation farther away. In contrast, agricultural and grassland areas were correlated with higher building density, higher fragmentation, and more dispersed building pattern possible legacies of agricultural settlement patterns. Understanding which factors influence building density and fragmentation is useful for landscape level planning and ecosystem management in northern Wisconsin and areas that share similar social and environmental constraints.     

Considering landscape connectivity and gene flow in the Anthropocene using complementary landscape genetics and habitat modelling approaches

Landscape Ecology - A comprehensive understanding of how rapidly changing environments affect species gene flow is critical for mitigating future biodiversity losses. While recent methodological...     

Intraspecific body size increases with habitat fragmentation in wild bee pollinators

Context

In modern agricultural landscapes, fragmentation of partial habitats is a significant filter for multi-habitat users, reducing local taxonomic and functional diversity. There is compelling evidence that small species are more susceptible than large species. The impact of habitat fragmentation on intraspecific body-size distribution, however, is yet unexplored.

Objectives

We tested habitat fragmentation, a major driver of pollinator loss, for its impact on intraspecific body-size distributions of solitary wild-bee species. Subsequently, we tested individual body size for its impact on pollination services.

Methods

We sampled 1272 individuals of the four most common Andrena wild bee species in 22 newly established flowering fields (0.21–0.41 ha) in Hessen, Central Germany, over two consecutive years. Study sites were located in a ca. 80 ha landscape context of increasing habitat fragmentation. We analysed the pollen loads of the most abundant species.

Results

Body size within local populations of the two medium-sized bees increased with fragmentation, suggesting intraspecific selection for higher dispersal capacity. Pollen analysis carried out for the most common species revealed that larger individuals visited a significantly smaller plant spectrum. Habitat fragmentation may thus alter pollination services without necessarily affecting species richness or composition.

Conclusions

Systematic body-size variation at the population level thus explains the considerable variability between simple community measures and ecosystem functioning. Filtering processes at the individual level require increased understanding for targeting pollination services under current and future land-use change.

     

Arthropod reaction to landscape and habitat features in agricultural landscapes

Determining explanatory environmental factors that lead to patterns of biodiversity in cultivated landscapes is an important step for the assessment of the impact of landscape changes. In the context of an assessment of the effect of agricultural national extensification programme on biodiversity, field data of 2 regions were collected according to a stratified sampling method. A distribution model of 3 indicator species taxa (butterflies, spiders, and carabid beetles) is related to influencing factors by means of multivariate statistics (CCA, partial CCA). Hypothetical influencing factors are categorised as follows: (1) habitat (habitat type, management techniques) and (2) landscape (habitat heterogeneity, variability, diversity, proportion of natural and semi-natural areas). The correlation models developed for spider, carabid beetle and butterfly assemblages revealed that there are no general rules relating species diversity to habitat and landscape features. The relationship strongly depends on the organism and on the region under study. Therefore, biodiversity response to landscape and habitat changes has to be identified by means of a multi-indicator concept in different landscape situations.This revised version was published online in May 2005 with corrections to the Cover Date.