Responses of Chilean forest birds to anthropogenic habitat fragmentation across spatial scales

Although it is recognized that anthropogenic forest fragmentation affects habitat use by organisms across multiple spatial scales, there is uncertainty about these effects. We used a hierarchical sampling design spanning three spatial scales of habitat variability (landscape > patch > within-patch) and generalized mixed-effect models to assess the scale-dependent responses of bird species to fragmentation in temperate forests of southern Chile. The abundances of nine of 20 bird species were affected by interactions across spatial scales. These interactions resulted in a limited effect of within-patch habitat structure on the abundance of birds in landscapes with low forest cover, suggesting that suitable local habitats, such as sites with dense understory cover or large trees, are underutilized or remain unused in highly fragmented landscapes. Habitat specialists and cavity-nesters, such as tree-trunk foragers and tapaculos, were most likely to exhibit interactions across spatial scales. Because providing additional sites with dense understory vegetation or large habitat trees does not compensate the negative effect of the loss of forest area on bird species, conservation strategies should ensure the retention of native forest patches in the mixed-use landscapes.     

Distribution of woodland amphibians along a forest fragmentation gradient

Understanding how changes in land-use affect the distribution and abundance of organisms is an increasingly important question in landscape ecology. Amphibians may be especially prone to local extinction resulting from human-caused transformation and fragmentation of their habitats owing to the spatially and temporally dynamic nature of their populations. In this study, distributions of five species of woodland amphibians with differing life histories were surveyed along a 10 km, spatially continuous gradient of forest fragmentation in southern Connecticut, U.S.A. Redback salamanders (Plethodon cinereus) and northern spring peepers (Pseudacris c. crucifer) occupied available habitat along the gradient's length. Wood frogs (Rana sylvatica) and spotted salamanders (Ambystoma maculatum) were absent from portions of the gradient where forest cover was reduced to below about 30%. Red-spotted newts (Notophthalmus v. viridescens) did not persist below a forest cover threshold of about 50%. Correlations between species' biological traits and their fragmentation tolerance imply that low density, population variability, and high mobility coupled with restricted habitat needs predispose woodland amphibians to local extinction caused by habitat fragmentation. These patterns are in contrast to the widely held notion that populations of the best dispersers are those most tolerant of habitat fragmentation.     

Using habitat extent and composition to predict the occurrence of woodland birds in fragmented landscapes

Landscape Ecology - The removal, alteration and fragmentation of habitat are key causes of biodiversity decline worldwide. In Australia, temperate woodlands have been disproportionately cleared...     

Deciduous forest and resident birds: the problem of fragmentation within a coniferous forest landscape

Six species of resident birds were censused in patches of deciduous forest within a coniferous forest landscape in south central Sweden. Here, the forests have been subjected to active forestry for a long time, but with recently increased intensity. Although the forest cover is more or less continuous in this landscape, mature deciduous forest is now a rare element compared with the untouched forest.All censused patches were similar with regards to size, proportion and amount of deciduous trees, but were either isolated in the coniferous forest (isolated patches) or near to other deciduous patches (aggregated patches). We concentrated on six species of resident birds, with moderate area requirements, that are tied to deciduous forest and whose ecology is well-known. The Nuthatch and the Marsh tit, which both show strict year-round territoriality and have a restricted dispersal phase, were significantly more likely to be found in aggregated than in isolated patches. No effect was found for the Great tit and the Blue tit, which are less territorial outside the breeding season and have a longer dispersal phase. Moreover, the Great tit is less specialized on deciduous forest than the other species. Also, the Long-tailed tit was negatively affected by isolation, which may be due to restricted dispersal and to larger area requirements of this flock-territorial species. The Hazel grouse, finally, was not affected, but this larger bird probably uses the forest in a different way from the smaller species.Our study clearly shows that fragmentation of one type of forest (deciduous) within another can have serious detrimental effects on forest-living species and raises important issues for forest management practices and conservation within a forest landscape.     

Modelling the effects of forest fragmentation on certain species of forest-breeding birds

Summary The influence of forest fragmentation was assessed on the abundance of six forest-breeding bird species. The study area (2327 sq Km) was located in south-west France. The forest cover, extracted from a Landsat MSS scene, was first reduced to a grid of 5865 quadrats, each 650 by 650 m. Two values were attributed with each quadrat: Quadrat Forest Cover (QFC), expressed in percent; and a local measure of forest fragmentation - the Neighbouring Forest Cover (NFC) - expressed on a 0–1000 scale. The distribution of six forest-breeding species was sampled on 556 quadrats.For each species, the local abundance appears to be more correlated with the fragmentation-oriented NFC value than with the local QFC value. For three species out of six (song thrush, robin, chaffinch) an incidence model, based on the Logistic regression, was built. A correct fit was obtained.An incidence map of these species was then built up over the whole study area. Their regional status was then estimated, for a sampling cost of less than 10% of censusing all the area.     

Sacrificing patches for linear habitat elements enhances metapopulation performance of woodland birds in fragmented landscapes

It is generally assumed that large patches of natural habitat are better for the survival of species than the same amount of habitat in smaller fragments or linear elements like hedges and tree rows. We use a spatially explicit individual-based model of a woodland bird to explore this hypothesis. We specifically ask whether mixtures of large, small and linear habitat elements are better for population performance than landscapes that consist of only large elements. With equal carrying capacity, metapopulations perform equally or better in heterogeneous landscape types that are a mix of linear, large and small habitat elements. We call this increased metapopulation performance of large and small elements “synergy”. These mixed conditions are superior because the small linear elements facilitate dispersal while patches secure the population in the long run because they have a lower extinction risk. The linear elements are able to catch and guide dispersing animals which results in higher connectivity between patches leading to higher metapopulation survival. Our results suggest that landscape designers should not always seek to conserve and create larger units but might better strive for more variable landscapes with mixtures of patch sizes and shapes. This is especially important when smaller units play a key role in connecting patches and dispersal through the matrix is poor.     

Effects of structural and functional habitat gaps on breeding woodland birds: working harder for less

The effects of habitat gaps on breeding success and parental daily energy expenditure (DEE) were investigated in great tits (Parus major) and blue tits (Cyanistes caeruleus) in urban parkland (Cardiff, UK) compared with birds in deciduous woodland (eastern England, UK). Tree canopy height, the percentage of gap in the canopy and the percentage of oak (in the wood only) within a 30 m radius of nest boxes were obtained from airborne remote-sensed data. Breeding success was monitored and parental DEE (great tits: both habitats; blue tits: park only) was measured using doubly labelled water in birds feeding young. In the park, mean (±SD) tree height (7.5 ± 4.7 m) was less than in the wood (10.6 ± 4.5 m), but the incidence of gaps (32.7 ± 22.6%) was greater (9.2 ± 14.7%). Great tits and blue tits both reared fewer young in the park and chick body mass was also reduced in park-reared great tits. Park great tits had a higher DEE (86.3 ± 12.3 kJ day⁻¹) than those in the wood (78.0 ± 11.7 kJ day⁻¹) and, because of smaller brood sizes, worked about 64% harder for each chick reared. Tits in the park with more than about 35% gap around their boxes had higher DEEs than the average for the habitat. In the wood, great tits with less oak around their boxes worked harder than average. Thus structural gaps, and functional gaps generated by variation in the quality of foraging habitat, increased the costs of rearing young. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Fine-scale effects of habitat loss and fragmentation despite large-scale gene flow for some regionally declining woodland bird species

Habitat loss and associated fragmentation effects are well-recognised threats to biodiversity. Loss of functional connectivity (mobility, gene flow and demographic continuity) could result in population decline in altered habitat, because smaller, isolated populations are more vulnerable to extinction. We tested whether substantial habitat reduction plus fragmentation is associated with reduced gene flow in three ??decliner?? woodland-dependent bird species (eastern yellow robin, weebill and spotted pardalote) identified in earlier work to have declined disproportionately in heavily fragmented landscapes in the Box-Ironbark forest region in north-central Victoria, Australia. For these three decliners, and one ??tolerant?? species (striated pardalote), we compared patterns of genetic diversity, relatedness, effective population size, sex-ratios and genic (allele frequency) differentiation among landscapes of different total tree cover, identified population subdivision at the regional scale, and explored fine-scale genotypic (individual-based genetic signature) structure. Unexpectedly high genetic connectivity across the study region was detected for ??decliner?? and ??tolerant?? species. Power analysis simulations suggest that moderate reductions in gene flow should have been detectable. However, there was evidence of local negative effects of reduced habitat extent and structural connectivity: slightly lower effective population sizes, lower genetic diversity, higher within-site relatedness and altered sex-ratios (for weebill and eastern yellow robin) in 10 × 10?km ??landscapes?? with low vegetation cover. We conclude that reduced structural connectivity in the Box-Ironbark ecosystem may still allow sufficient gene flow to avoid the harmful effects of inbreeding in our study species. Although there may still be negative consequences of fragmentation for demographic connectivity, the high genetic connectivity of mobile bird species in this system suggests that reconnecting isolated habitat patches may be less important than increasing habitat extent and/or quality if these need to be traded off.     

Sensitivity to habitat fragmentation across European landscapes in three temperate forest herbs

Landscape Ecology - Evidence for effects of habitat loss and fragmentation on the viability of temperate forest herb populations in agricultural landscapes is so far based on population genetic...     

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.     

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.     

Boundary effects on dispersal between habitat patches by forest birds (Parus major,P. caeruleus)

The behaviour of individuals in response to patch boundaries is acrucial element in many dispersal models. Diffusion-based models of dispersalpredict that both the likelihood and direction of dispersal from a habitatpatchare influenced by the starting position of a disperser in relation to the patchboundary. An alternative view is that the decision to disperse between patchesis uncoupled from movements within the patch of departure. The latter situationis most likely in the case of relatively mobile animals living in small patcheswith strongly reflecting boundaries. I tested the relationship betweenproximityto a boundary and natal dispersal in Great and Blue Tits born in relativelysmall (7-11 ha) forest patches with high population density innorthern Belgium. Birdsthat were born closer to the forest edge were not more likely to be recruitedoutside than inside the natal patch. However, Great Tits showed a significanttendency to emigrate in the direction of the nearest patch border. No sucheffect was found in Blue Tits. A possible explanation is that in Great Tits thedirection of dispersal, but not the decision to emigrate, is influenced by aprocess of familiarization with the area around the natal territory, includingareas across the patch border.This revised version was published online in May 2005 with corrections to the Cover Date.     

Habitat fragmentation and parasitism of a forest damselfly

We compared populations of a forest damselfly —Calopteryx maculata — in two kinds of landscapes. In fragmented landscapes, forested foraging patches were separated from streams (where oviposition and mating occur) by up to 500 m of pasture. In non-fragmented landscapes, there was continuous forest cover adjacent to streams. The prevalence and intensity of midgut infections of a gregarine parasite were significantly lower in the fragmented landscapes than in the non-fragmented landscapes. We have shown elsewhere that in the fragmented landscapes, damselflies move over greater areas to forage than in the non-fragmented landscapes. We postulate that these movements lower the rate of encounter between damselflies and oocysts, thus lowering the prevalence and intensity of infection. The differences suggest that actual habitat fragmentation events would alter the relationship between host and parasite, but that populations of both species would persist after fragmentation. Prevalence of parasitism is related to age but we found no residual effects of size on parasitism.     

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.     

Effects of land use legacies and habitat fragmentation on salamander abundance

Context

Landscape modification is an important driver of biodiversity declines, yet we lack insight into how ongoing landscape change and legacies of historical land use together shape biodiversity.

Objectives

We examined how a history of agricultural land use and current forest fragmentation influence the abundance of red-backed salamanders (Plethodon cinereus). We hypothesized that historical agriculture and fragmentation cause changes in habitat quality and landscape structure that limit abundance.

Methods

We measured salamander abundance at 95 forested sites in New York, USA, and we determined whether sites were agricultural fields within the last five decades. We used a structural equation model to estimate relationships between historical agriculture and salamander abundance mediated by changes in forest vegetation, microclimate, and landscape structure.

Results

Historical agriculture affected salamander abundance by altering forest vegetation at a local scale and forest cover at a landscape scale. Abundance was lowest at post-agricultural sites with low woody vegetation, leaf litter depth, and canopy cover. Post-agricultural sites had limited forest cover in the surrounding landscape historically, and salamander abundance was positively related to historical forest cover, suggesting that connectivity to source populations affects colonization of regenerating forests. Abundance was also negatively related to current forest fragmentation.

Conclusions

Historical land use can have legacy effects on animal abundance on par with effects of ongoing landscape change. We showed that associations between animal abundance and historical land use can be driven by altered site conditions and surrounding habitat area, indicating that restoration efforts should consider local site conditions and landscape context.

     

Influence of high-resolution data on the assessment of forest fragmentation

Context

Remote sensing has been a foundation of landscape ecology. The spatial resolution (pixel size) of remotely sensed land cover products has improved since the introduction of landscape ecology in the United States. Because patterns depend on spatial resolution, emerging improvements in the spatial resolution of land cover may lead to new insights about the scaling of landscape patterns.

Objective

We compared forest fragmentation measures derived from very high resolution (1 m²) data with the same measures derived from the commonly used (30 m?×??30 m; 900 m²) Landsat-based data.

Methods

We applied area-density scaling to binary (forest; non-forest) maps for both sources to derive source-specific estimates of dominant (density ≥?60%), interior (≥?90%), and intact (100%) forest.

Results

Switching from low- to high-resolution data produced statistical and geographic shifts in forest spatial patterns. Forest and non-forest features that were “invisible” at low resolution but identifiable at high resolution resulted in higher estimates of dominant and interior forest but lower estimates of intact forest from the high-resolution source. Overall, the high-resolution data detected more forest that was more contagiously distributed even at larger spatial scales.

Conclusion

We anticipate that improvements in the spatial resolution of remotely sensed land cover products will advance landscape ecology through re-interpretations of patterns and scaling, by fostering new landscape pattern measurements, and by testing new spatial pattern-ecological process hypotheses.

     

The landscape matrix modifies the effect of habitat fragmentation in grassland butterflies

The landscape matrix is suggested to influence the effect of habitat fragmentation on species richness, but the generality of this prediction has not been tested. Here, we used data from 10 independent studies on butterfly species richness, where the matrix surrounding grassland patches was dominated by either forest or arable land to test if matrix land use influenced the response of species richness to patch area and connectivity. To account for the possibility that some of the observed species use the matrix as their main or complementary habitat, we analysed the effects on total species richness and on the richness of grassland specialist and non-specialist (generalists and specialists on other habitat types) butterflies separately. Specialists and non-specialists were defined separately for each dataset. Total species richness and the richness of grassland specialist butterflies were positively related to patch area and forest cover in the matrix, and negatively to patch isolation. The strength of the species-area relationship was modified by matrix land use and had a slope that decreased with increasing forest cover in the matrix. Potential mechanisms for the weaker effect of grassland fragmentation in forest-dominated landscapes are (1) that the forest matrix is more heterogeneous and contains more resources, (2) that small grassland patches in a matrix dominated by arable land suffer more from negative edge effects or (3) that the arable matrix constitutes a stronger barrier to dispersal between populations. Regardless of the mechanisms, our results show that there are general effects of matrix land use across landscapes and regions, and that landscape management that increases matrix quality can be a complement to habitat restoration and re-creation in fragmented landscapes.     

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.     

Scale-dependent habitat use in three species of prairie wetland birds

We evaluated the influence of scale on habitat use for three wetland-obligate bird species with divergent life history characteristics and possible scale-dependent criteria for nesting and foraging in South Dakota, USA. A stratified, two-stage cluster sample was used to randomly select survey wetlands within strata defined by region, wetland density, and wetland surface area. We used 18-m (0.1 ha) fixed radius circular-plots to survey birds in 412 semipermanent wetlands during the summers of 1995 and 1996. Variation in habitat use by pied-billed grebes (Podilymbus podiceps) and yellow-headed blackbirds (Xanthocephalus xanthocephalus), two sedentary species that rarely exploit resources outside the vicinity of nest wetlands, was explained solely by within-patch variation. Yellow-headed blackbirds were a cosmopolitan species that commonly nested in small wetlands, whereas pied-billed grebes were an area-sensitive species that used larger wetlands regardless of landscape pattern. Area requirements for black terns (Chlidonias niger), a vagile species that typically forages up to 4 km away from the nest wetland, fluctuated in response to landscape structure. Black tern area requirements were small (6.5 ha) in heterogeneous landscapes compared to those in homogeneous landscapes (15.4–32.6 ha). Low wetland density landscapes composed of small wetlands, where few nesting wetlands occurred and potential food sources were spread over large distances, were not widely used by black terns. Landscape-level measurements related to black tern occurrence extended past relationships between wetlands into the surrounding matrix. Black terns were more likely to occur in landscapes where grasslands had not been tilled for agricultural production. Our findings represent empirical evidence that characteristics of entire landscapes, rather than individual patches, must be quantified to assess habitat suitability for wide-ranging species that use resources over large areas.     

Confronting collinearity: comparing methods for disentangling the effects of habitat loss and fragmentation

Estimating the relative importance of habitat loss and fragmentation is necessary to estimate the potential benefits of specific management actions and to ensure that limited conservation resources are used efficiently. However, estimating relative effects is complicated because the two processes are highly correlated. Previous studies have used a wide variety of statistical methods to separate their effects and we speculated that the published results may have been influenced by the methods used. We used simulations to determine whether, under identical conditions, the following 7 methods generate different estimates of relative importance for realistically correlated landscape predictors: residual regression, model or variable selection, averaged coefficients from all supported models, summed Akaike weights, classical variance partitioning, hierarchical variance partitioning, and a multiple regression model with no adjustments for collinearity. We found that different methods generated different rankings of the predictors and that some metrics were strongly biased. Residual regression and variance partitioning were highly biased by correlations among predictors and the bias depended on the direction of a predictor’s effect (positive vs. negative). Our results suggest that many efforts to deal with the correlation between amount and fragmentation may have done more harm than good. If confounding effects are controlled and adequate thought is given to the ecological mechanisms behind modeled predictors, then standardized partial regression coefficients are unbiased estimates of the relative importance of amount and fragmentation, even when predictors are highly correlated.