Effects of fragmentation on functional diversity associated with aboveground biomass in a high Andean forest in Colombia

Context

Forest fragmentation alters the composition, structure and function of ecosystems and affects ecological processes that are fundamental for the provision of ecosystem services where functional diversity is sensitive to its effects. Analyzing the functional responses of the plant community to fragmentation can provide new approaches to its conservation and management.

Objectives

We analyzed whether the functional diversity of woody individuals associated with aboveground biomass (AGB) in a high Andean forest in Colombia is affected by fragmentation.

Methods

Based on three fragmentation categories identified using landscape metrics, we selected ten forest fragments. Multitrait and monotrait functional diversity indexes (foliar and wood) weighted by aboveground biomass were calculated in plots of 0.1 ha in each fragment. Analysis of variance was performed, and simple linear regressions were quantified to identify the relationships between functional diversity and fragmentation.

Results

The category of large fragments had a higher average AGB than did the medium and small fragments. Fragmentation had effects on the variance of some foliar and stem traits but not on functional dominance. For the multitraits indexes, the edge contrast was negatively related with functional dispersion.

Conclusions

The categories analyzed have similar responses in terms of functionality associated with AGB. We highlight the importance of small fragments in the maintenance of plant functional diversity and as reservoirs of AGB. We underline that small fragments are important to consider in the development of conservation and connectivity strategies.

     

Correlations and variance among species traits explain contrasting impacts of fragmentation and habitat loss on functional diversity

Landscape Ecology - Understanding how landscape fragmentation affects functional diversity, defined as the distribution of functional traits in an assemblage, is critical for managing landscapes...     

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.     

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.     

Habitat alteration and habitat fragmentation differentially affect beta diversity of stream fish communities

Context

The cumulative impact of broad scale environmental change includes altered land-cover and fragmentation. Both altered land-cover and fragmentation have a negative effect on species diversity, but the scale they act on may differ because land-cover alters environmental characteristics, whereas fragmentation alters movement among sites.

Objectives

We evaluated the scale specific effects of land-cover, fragmentation, and habitat size on alpha and beta diversity (total, turnover, and nestedness).

Methods

Stream fish communities were sampled across five urbanizing watersheds. Generalized mixed linear models were used to test how diversity (alpha and beta) is affected by land-cover, connectivity, and habitat size. Indices of land-cover were calculated from correspondence analyses on land-cover data, fragmentation was estimated with the dendritic connectivity index, and habitat size was calculated as the length of the stream segment (alpha diversity) or the length of the stream network (beta diversity).

Results

Alpha diversity was most strongly related to land-cover variables associated with urban development and agriculture (negative relationship with urbanization). Whereas, beta diversity was most strongly influenced by habitat size (positive relationship) and fragmentation (positive relationship). Turnover was positively correlated with fragmentation and habitat size, whereas species loss was negatively correlated with habitat size.

Conclusions

Land-cover has a larger effect on alpha diversity because it alters the environmental conditions at a site, whereas fragmentation has a larger effect on beta diversity because it affects the movement of individuals among sites. Assessing the cumulative impact of environmental change requires a multiscale approach that simultaneously considers alpha and beta diversity.

     

Loss and fragmentation of mature woodland reduce the habitat niche breadth of forest birds

Context

Habitat loss and fragmentation may alter habitat occupancy patterns, for example through a reduction in regional abundance or in functional connectivity, which in turn may reduce the number of dispersers or their ability to prospect for territories. Yet, the relationship between landscape structure and habitat niche remains poorly known.

Objectives

We hypothesized that changes in landscape structure associated with habitat loss and fragmentation will reduce the habitat niche breadth of forest birds, either through a reduction in density-dependent spillover from optimal habitat or by impeding the colonization of patches.

Methods

We surveyed forest birds with point counts in eastern Ontario, Canada, and analyzed their response to loss and fragmentation of mature woodland. We selected 62 landscapes varying in both forest cover (15–45%) and its degree of fragmentation, and classified them into two categories (high versus low levels of loss and fragmentation). We determined the habitat niche breadth of 12 focal species as a function of 8 habitat structure variables for each landscape category.

Results

Habitat niche breadth was narrower in landscapes with high versus low levels of loss and fragmentation of forest cover. The relative occupancy of marginal habitat appeared to drive this relationship. Species sensitivity to mature forest cover had no apparent influence on relative niche breadth.

Conclusions

Regional abundance and, in turn, density-dependent spillover into suboptimal habitat appeared to be determinants of habitat niche breadth. For a given proportion of forest cover, fragmentation also appeared to alter habitat use, which could exacerbate its other negative effects unless functional connectivity is high enough to allow individuals to saturate optimal habitat.

     

Effects of spatial habitat heterogeneity on habitat selection and annual fecundity for a migratory forest songbird

Understanding how spatial habitat patterns influence abundance and dynamics of animal populations is a primary goal in landscape ecology. We used an information-theoretic approach to investigate the association between habitat patterns at multiple spatial scales and demographic patterns for black-throated blue warblers (Dendroica caerulescens) at 20 study sites in west-central Vermont, USA from 2002 to 2005. Sites were characterized by: (1) territory-scale shrub density, (2) patch-scale shrub density occurring within 25 ha of territories, and (3) landscape-scale habitat patterns occurring within 5 km radius extents of territories. We considered multiple population parameters including abundance, age ratios, and annual fecundity. Territory-scale shrub density was most important for determining abundance and age ratios, but landscape-scale habitat structure strongly influenced reproductive output. Sites with higher territory-scale shrub density had higher abundance, and were more likely to be occupied by older, more experienced individuals compared to sites with lower shrub density. However, annual fecundity was higher on sites located in contiguously forested landscapes where shrub density was lower than the fragmented sites. Further, effects of habitat pattern at one spatial scale depended on habitat conditions at different scales. For example, abundance increased with increasing territory-scale shrub density, but this effect was much stronger in fragmented landscapes than in contiguously forested landscapes. These results suggest that habitat pattern at different spatial scales affect demographic parameters in different ways, and that effects of habitat patterns at one spatial scale depends on habitat conditions at other scales.     

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.     

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.     

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.     

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.     

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.     

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...     

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.

     

Separating the effects of habitat amount and fragmentation on invertebrate abundance using a multi-scale framework

Context

Herbicide treatments in viticulture can generate highly contrasting mosaics of vegetated and bare vineyards, of which vegetated fields often provide better conditions for biodiversity. In southern Switzerland, where herbicides are applied at large scales, vegetated vineyards are limited in extent and isolated from one another, potentially limiting the distribution and dispersal ability of organisms.

Objectives

We tested the separate and interactive effects of habitat amount and fragmentation on invertebrate abundance using a multi-scale framework, along with additional environmental factors. We identified which variables at which scales were most important in predicting patterns of invertebrate abundance.

Methods

We used a factorial design to sample across a gradient of habitat amount (area of vegetated vineyards, measured as percentage of landscape PLAND) and fragmentation (number of vegetated patches, measured as patch density PD). Using 10 different spatial scales, we identified the factors and scales that most strongly predicted invertebrate abundance and tested potential interactions between habitat amount and fragmentation.

Results

Habitat amount (PLAND index) was most important in predicting invertebrate numbers at a field scale (50 m radius). In contrast, we found a negative effect of fragmentation (PD) at a broad scale of 450 m radius, but no interactive effect between the two.

Conclusions

The spatial scales at which habitat amount and fragmentation affect invertebrates differ, underpinning the importance of spatially explicit study designs in disentangling the effects between habitat amount and configuration. We showed that the amount of vegetated vineyards has more influence on invertebrate abundance, but that fragmentation also contributed substantially. This suggests that efforts for augmenting the area of vegetated vineyards is more beneficial for invertebrate numbers than attempts to connect them.

     

Spatial and temporal variability of fragmentation effects in a long term,eucalypt forest fragmentation experiment

Context

Although forest fragmentation is generally thought to impact tree growth and mortality negatively, recent work suggests some forests are resilient. Experimental forests provide an opportunity to examine the timing and extent of forest tree resilience to disturbance from fragmentation.

Objectives

We used the Wog Wog Habitat Fragmentation Experiment in southeastern Australia to test Eucalyptus growth and survivorship responses to forest fragmentation over a 26 year period.

Methods

We measured 2418 tree diameters and used spline-regression techniques to examine non-monotonic fragmentation effect over two time periods.

Results

Over the first 4 years after fragmentation, individual eucalypt tree growth was greater than in continuous forest for large trees and mortality rates were higher only within 10 m of edges. Over the following 22 years only the effects on tree growth remained and on average all fragments rebounded so that their biomass and mortality rates were equivalent to continuous forest. Importantly non-monotonic patterns were observed in growth and mortality with respect to area and distance from edge in both study periods, demonstrating that fragmentation impacts on trees can be strong in localized areas (greatest in 3 ha fragments and 0–30 m edges) and over short time periods.

Conclusions

Dry-sclerophyll eucalypt forests join the set of forest types that display resilient growth dynamics post fragmentation. Moreover, persistent non-monotonic impacts on tree growth with respect to tree size, fragment area, and fragment distance from edge, highlighting landscape fragmentation as a driver of habitat heterogeneity within remnant forest fragments.

     

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.     

Effect of disturbances and habitat fragmentation on an arboreal habitat specialist mammal using GPS telemetry: a case of the red panda

Landscape Ecology - Habitat specialists residing in human-modified landscapes are likely to be more vulnerable to disturbance because of a functional reliance on very particular habitat features....     

Metapopulation theory and habitat fragmentation: a review of holarctic breeding bird studies

Metapopulations are conceived as spatially structured populations consisting of distinct units (subpopulations), separated by space or barriers, and connected by dispersal movements. Metapopulations characteristically demonstrate a turnover of local populations going extinct and becoming re-established, resulting in a distribution pattern that shifts over time. Metapopulation theory is used to analyse the effects of habitat fragmentation on birds in the temperate zone, integrating various explanations for the paucity of species in isolated ecotopes.There is some evidence that turnover of local populations occurs in fragmented systems. A few studies based on time series demonstrate the local extinction rate to be related to the size of the habitat fragment, whereas the recolonization rate depends on the degree of isolation. Most evidence comes from short-term pattern studies in which the probability of occurrence was found to depend on the size of habitat fragments, on their relative position in the landscape and on the density of corridors lowering the landscape resistance. These data are consistent with predictions from metapopulation theory. However, almost all investigations consider wood fragmentation in agricultural landscapes, and there is a great need for studies in naturally fragmented landscapes as well as for studies focussing on other, less predictable, habitat types.     

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.