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1.

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.
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2.

Context

The positive correlation between landscape area of semi-natural habitat and wild pollinator richness and abundance in agroecosystems has been well studied. However, we lack a deep understanding of local scale floral resource and nest provisioning for wild bees necessary to optimize implementation of pollinator conservation practices.

Objectives

The primary objective of this study was to use a spatially interactive landscape pollination model (hereafter, the Lonsdorf model) to represent field scale spatial patterns of wild bee abundance and richness within a heterogeneous landscape in the mid-Atlantic USA.

Methods

We parameterized the Lonsdorf model with high resolution aerial imagery and insight from a previously published floristic study. To test the Lonsdorf model predictions, field studies were conducted to measure wild bee abundance and species richness in apple orchards as a function of distance from a forest edge.

Results

Field measurements indicated apple pollinator abundance and species richness significantly decreased with increasing distance from the forest edge. The Lonsdorf model pollination service score was highly sensitive to changes in resource provisioning in orchard and non-crop areas, and including resource rich forest and forest edge habitats in the model significantly improved pollination service estimates.

Conclusions

We demonstrated a novel application of the Lonsdorf model at a field scale to predict trends in pollination service provisioning as a factor of local habitat features. With sufficiently detailed inputs, the Lonsdorf model is a promising tool to quantify field scale pollination service deficits, guiding more cost effective habitat supplementation and other conservation efforts.
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3.

Context

The relationship between biodiversity and ecosystem functioning (BEF) has been a central topic in ecology for more than 20 years. While experimental and theoretical studies have produced much knowledge of how biodiversity affects ecosystem functioning, it remains poorly understood how habitat fragmentation affects the BEF relationship.

Objectives

To develop a framework that connects habitat fragmentation to the BEF relationship from a landscape perspective.

Methods

We reviewed the literature on habitat fragmentation, BEF, and related fields, and developed a framework to analyze how habitat fragmentation affects the BEF relationship through altering biodiversity, environmental conditions, and both, based on the pattern-process-scale perspective in landscape ecology.

Results

Our synthesis of the literature suggests that habitat fragmentation can alter BEF relationship through several processes. First, habitat fragmentation causes the non-random loss of species that make major contributions to ecosystem functioning (decreasing sampling effect), and reduces mutualistic interactions (decreasing complementarity effects) regardless of the changes in species richness. Second, environmental conditions within patches and ecological flows among patches vary significantly with the degree of fragmentation, which potentially contributes to and modulates the BEF relationship.

Conclusions

Habitat fragmentation can affect the BEF relationship directly by altering community composition, as well as indirectly by changing environmental conditions within and among habitat patches on both local and landscape levels. The BEF relationship obtained from small plots and over short time periods may not fully represent that in real landscapes that are fragmented, dynamic, and continuously influenced by myriad human activities on different scales in time and space.
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4.

Context

The umbrella approach applied to landscape connectivity is based on the principle that the conservation or restoration of the dispersal habitats for some species also can facilitate the movement of others. Species traits alone do not seem to be enough to identify good connectivity umbrella species, showing the need to investigate the influence of additional factors on this property.

Objectives

We test whether the potential of a species as a connectivity umbrella can be influenced by landscape composition and configuration.

Methods

We simulated movement routes for eight hypothetical species in artificial patchy landscapes with different levels of fragmentation, habitat amount and matrix permeability. We determined the effectiveness of the connectivity umbrella of the virtual species using pairwise intersections of important habitats for their movements in all landscapes.

Results

The connectivity umbrella performance of all species was affected by the interaction of fragmentation level and habitat amount. In general, species performance increased with decreasing fragmentation and increasing habitat amount. In most landscapes and considering the same dispersal threshold, species able to move more easily through the matrix showed higher umbrella performance than those for which the matrix offered greater resistance.

Conclusions

The connectivity umbrella is not a static feature that depends only on the species traits, but rather a dynamic property that also varies according to the landscape attributes. Therefore, we do not recommend spatial transferability of the connectivity umbrella species identified in a landscape to others that have divergent levels of fragmentation and habitat quantity.
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5.

Context

Native vegetation is often used as a proxy for habitat to estimate habitat availability in landscapes. This approach may lead to incorrect estimates of the impacts of habitat loss and fragmentation on species, which have not been thoroughly quantified so far.

Objectives

We quantified to what extent the loss of native vegetation reflect actual habitat loss by native species in landscapes. We tested the hypothesis that habitat availability declines at greater rates than native vegetation and thus is overestimated when it is quantified on the basis of native vegetation.

Methods

Using simulations, we quantified how the loss of native vegetation in artificial and real landscapes affects habitat availability for species with different habitat requirements. We contrasted a generalist species, which uses all native vegetation, with 10 habitat-specialist species classified into three categories (interior, patchy and riparian species).

Results

Habitat availability generally declined at greater rates than native vegetation for all specialist species. This pattern was apparent for different specialist species in a broad range of landscape types. Interior species always lost habitat availability more rapidly than the generalist species. Most riparian species lost habitat availability more rapidly than the generalist species. Responses of patchy species were more complex, depending on their dispersal abilities and landscape structure.

Conclusions

Habitat availability is likely to be overestimated when native vegetation is used as proxy for habitat, because habitat availability will generally decline at greater rates than native vegetation. Therefore, a species-centered approach should be adopted when estimating habitat availability in landscapes.
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6.

Context

Habitat loss and habitat fragmentation negatively affect amphibian populations. Roads impact amphibian species through barrier effects and traffic mortality. The landscape variable ‘accessible habitat’ considers the combined effects of habitat loss and roads on populations.

Objectives

The aim was to test whether accessible habitat was a better predictor of amphibian species richness than separate measures of road effects and habitat loss. I assessed how accessible habitat and local habitat variables determine species richness and community composition.

Methods

Frog and tadpole surveys were conducted at 52 wetlands in a peri-urban area of eastern Australia. Accessible habitat was delineated using a highway. Regressions were used to examine relationships between species richness and eleven landscape and local habitat variables. Redundancy analysis was used to examine relationships between community composition and accessible habitat and local habitat variables.

Results

Best-ranked models of species richness included both landscape and local habitat variables. There were positive relationships between species richness and accessible habitat and distance to the highway, and uncertain relationships with proportion cover of native vegetation and road density. There were negative relationships between species richness and concreted wetlands and wetland electrical conductivity. Four species were positively associated with accessible habitat, whereas all species were negatively associated with wetland type.

Conclusions

Barrier effects caused by the highway and habitat loss have negatively affected the amphibian community. Local habitat variables had strong relationships with species richness and community composition, highlighting the importance of both availability and quality of habitat for amphibian conservation near major roads.
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7.

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.
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8.

Context

Landscape changes can be an important modifier of disease. Habitat fragmentation commonly results in reduced connectivity in host populations and increased use of the remaining habitat. For environmentally transmitted parasites, this presents a possible trade-off between transmission potential at the local and global level.

Objectives

We quantify the effects of fragmentation on the transmission of an environmentally transmitted parasite, teasing apart the relative effects of habitat composition and configuration on both host movement behaviour and subsequent infection patterns.

Methods

We use a spatially-explicit epidemiological model to simulate the effects of habitat fragmentation, using, as an example, whipworm (Trichuris sp.) within a red colobus monkey population (Procolobus rufomitratus).

Results

We found that habitat fragmentation did not always lead to a trade-off between population connectivity and concentration of habitat use in host movement behaviour or in final population infection patterns. However, our simulation results suggest the spatial configuration of the remaining habitat became increasingly influential on behavioural and infection outcomes as habitat was removed. Additionally, we found common fragmentation metrics provided little ability to explain variation in propagation of infections.

Conclusions

Our results suggest an interaction between habitat configuration and composition should be considered when assessing disease related impacts of habitat fragmentation on environmentally transmitted parasites, especially in cases where habitat loss is high (≥?30%). We also propose that spatially-explicit simulations that capture a host’s response to fragmentation could aid in the development of novel landscape metrics targeted towards specific host-parasite-landscape systems.
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9.

Context

Distribution and connectivity of suitable habitat for species of conservation concern is critical for effective conservation planning. Capercaillie (Tetrao urogallus), an umbrella species for biodiversity conservation, is increasingly threatened because of habitat loss and fragmentation.

Objective

We assessed the impact of drastic changes in forest management in the Carpathian Mountains, a major stronghold of capercaillie in Europe, on habitat distribution and connectivity.

Methods

We used field data surveys with a forest disturbance dataset for 1985–2010 to map habitat suitability, and we used graph theory to analyse habitat connectivity.

Results

Climate, topography, forest proportion and fragmentation, and the distance to roads and settlements best identified capercaillie presence. Suitable habitat area was 7510 km2 in 1985; by 2010, clear-cutting had reduced that area by 1110 km2. More suitable habitat was lost inside protected areas (571 km2) than outside (413 km2). Habitat loss of 15 % reduced functional connectivity by 33 % since 1985.

Conclusions

Forest management, particularly large-scale clear-cutting and salvage logging, have substantially diminished and fragmented suitable capercaillie habitat, regardless of the status of forest protection. Consequently, larger areas with suitable habitat are now isolated and many patches are too small to sustain viable populations. Given that protection of capercaillie habitat would benefit many other species, including old-growth specialists and large carnivores, conservation actions to halt the loss of capercaillie habitat is urgently needed. We recommend adopting policies to protect natural forests, limiting large-scale clear-cutting and salvage logging, implementing ecological forestry, and restricting road building to reduce forest fragmentation.
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10.

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.
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11.

Context

The anthropocene is characterised by global landscape modification, and the structure of remnant habitats can explain different patterns of species richness. The most pervasive processes of degradation include habitat loss and fragmentation. However, a recovery of modified landscape is occurring in some areas.

Objectives

The main goal is to know how lichen and bryophyte epiphytic richness growing on Mediterranean forests is influenced not only by fragments characteristics but also by the structure of the landscape. We introduce a temporal dimension in order to evaluate if the historical landscape structure is relevant for current epiphytic communities.

Methods

40 well-preserved forest fragments were selected in a landscape with a large habitat loss over decades, but with a recovery of forest surface in the last 55 years. The most relevant fragment and landscape-scale attributes were considered. Some of the variables were measured in three different years to incorporate a temporal framework.

Results

The results showed that variables at fragment scale had a higher influence, whereas variables at the landscape scale were irrelevant. Among all the historical variables analyzed, only the shift in forest fragment size had influence on species richness.

Conclusions

Mediterranean forests had suffered fragmentation along centuries. Their epiphytic communities also suffer the hard conditions of Mediterranean climate. Our results indicate that Mediterranean epiphytic communities may be in a threshold since it they will never be similar to those communities existing previous fragmentation process even a recovery habitat occur or, they may require more time to response to this habitat recovery.
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12.

Context

Understanding the factors contributing to maintaining biodiversity is crucial to mitigate the impact of anthropogenic disturbances. Representing large proportions of green area in highly modified landscapes, residential gardens are often seen as local habitats that can contribute to larger networks of suitable environments at the landscape scale.

Objectives

We investigated the impact of the landscape context on butterfly communities observed in residential gardens, taking into account garden characteristics, land-use types and presence of linear features in the surrounding landscape. We examined how species traits affected butterflies’ response to landscape context and habitat quality.

Methods

We performed a cross-scale study, based on citizen science data documenting butterfly species composition and abundance in 920 gardens across France. We examined the effect of garden quality, the area of different land-use types and the length of linear elements measured at three scales within the surrounding landscape. Species were grouped according to their habitat preference and mobility.

Results

Urbanization negatively affected total species richness and the abundance of butterfly in each group. This was related to declining habitat quality and reduced area of suitable habitat in the surrounding landscape. The magnitude of this effect, however, was negatively correlated with mobility, a trait related to habitat preference. The spatial scale at which landscape context best explained variation in butterfly abundance changed with species’ habitat preference.

Conclusions

This study highlights the importance of preserving high quality habitats in altered landscapes and considering species’ mobility and habitat preference when assessing the impact of landscapes on butterfly communities.
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13.

Context

Species show different sensitivity to habitat loss and fragmentation depending on their specialization. Populations of a species at the range margin are generally assumed to be more stenoecious than populations at the core of the distribution and should therefore be more sensitive to habitat fragmentation.

Objectives

We evaluated the hypothesis that fragmentation effects species more strongly at the range periphery of their range compared to the core, resulting in lower genetic variability in comparable patch sizes and lower gene flow among populations.

Methods

We compared the genetic diversity and structure of five sand lizard (Lacerta agilis) populations at the margin of its range in Bulgaria and of 11 populations at the core of its distribution in Germany. We based the analysis on microsatellites, comprising 15 loci in Bulgaria and 12 in Germany.

Results

All diversity indices declined with patch size. For medium-sized patches all diversity indices were lower at the range periphery compared to the core, with two of them being significant. AICc based model selection showed strong support for core/periphery and patch size effects for observed and expected heterozygosity but only a patch size effect for allelic richness. There was no isolation-by-distance and each sampled population was allocated to a separate cluster with high probability for both countries, indicating that all populations are (almost) completely isolated.

Conclusion

Our study indicates an increased sensitivity of a species to fragmentation at the periphery compared to the core of its distribution. This differential sensitivity should be accounted for when prioritizing species based on their fragmentation sensitivity in landscape management.
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14.

Context

Land use changes have modified the extent and structure of native vegetation, resulting in fragmentation of native species habitat. Connectivity is increasingly seen as a requirement for effective conservation in these landscapes, but the question remains: ‘connectivity for which species?’.

Objective

The aim of this study was to develop and then apply a rapid, expert-based, dispersal guild approach where species are grouped on similar fine-scale dispersal behaviour (such as between scattered trees) and habitat characteristics.

Methods

Dispersal guilds were identified using clustering techniques to compare dispersal and habitat parameters elicited from experts. We modelled least-cost paths and corridors between patches and individual movement probabilities within these corridors for each of the dispersal guilds using Circuitscape. We demonstrate our approach with a case study in the Tasmanian Northern Midlands, Australia.

Results

The dispersal guild approach grouped the 12 species into five dispersal guilds. The connectivity modelling of those five guilds found that broadly dispersing species in this landscape, such as medium-sized carnivorous mammals, were unaffected by fragmentation while from the perspective of the three dispersal guilds made up of smaller mammals, the landscape appeared highly fragmented.

Conclusions

Our approach yields biologically defensible outputs that are broadly applicable, particularly for conservation planning where data and resources are limited. It is a useful first step in multi-species conservation planning which aims to identify those species most in need of conservation efforts.
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15.

Context

The local intensity of farming practices is considered as an important driver of biodiversity in agricultural landscapes and its effect on biodiversity has been shown to interact with landscape complexity. But the influence of landscape-wide intensity of farming practices on biodiversity and its combined effect with landscape complexity have been little explored.

Objective

In this study, we tested the interactive effect of the landscape-wide intensity of farming practices and landscape complexity on the local species richness and abundance of farmland wild bee communities.

Methods

We captured wild bees in 96 crop fields and explored the effect of landscape-wide intensity of various farming practices along a gradient of landscape complexity (proportion of semi-natural habitats).

Results

We found that species richness and abundance of wild bees were more positively influenced by landscape complexity in highly insecticide-sprayed landscapes than in less intensively managed landscapes. In contrast, we found that the positive effect of landscape complexity on bee species richness only occurred in landscapes with low nitrogen inputs.

Conclusions

Our study demonstrates the interactive effects of landscape-wide farming intensity and landscape complexity in shaping the diversity of farmland wild bee communities. We conclude that the management of farming intensity at the landscape-scale could mitigate the effects of habitat loss on wild bee decline and would help to maintain pollination services in agricultural landscapes.
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16.

Context

Habitat loss is a major threat to biodiversity. It can create temporal lags in decline of species in relation to destruction of habitat coverage. Plant species specialized in semi-natural grasslands, especially meadows, often express such extinction debt.

Objectives

We studied habitat loss and fragmentation of meadows and examined whether the changes in meadow coverage had caused an extinction debt on vascular plants. We also studied whether historical or present landscape patterns or contemporary environmental factors were more important determinants of species occurrence.

Methods

We surveyed the plant species assemblages of 12 grazed and 12 mown meadows in Central Finland and detected the meadow coverages from their surroundings on two spatial scales and on three time steps. We modelled the effects of functional connectivity, habitat amount, and isolation on species richness and community composition.

Results

We observed drastic and dynamic meadow loss in landscapes surrounding our study sites during the last 150 years. However, we did not find explicit evidence for an extinction debt in meadow plants. The observed species richness correlated with contemporary factors, whereas both contemporary factors and habitat availability during the 1960s affected community composition.

Conclusions

Effective conservation management of meadow biodiversity builds on accurate understanding of the relative importance of past and present factors on species assemblages. Both mown and grazed meadows with high species richness need to be managed in the future. The management effort should preferably be targeted to sites located near to each other.
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17.

Context

Increasing human populations in urban areas pose a threat to species’ persistence through habitat loss and fragmentation. It is therefore essential that we develop methods to investigate critical habitat loss thresholds and least detrimental landscape configurations.

Objectives

We develop a framework to assess how the pattern of habitat loss impacts the ecological and social characteristics of a landscape and how this varies depending on the species and criteria by which it is judged.

Methods

We use a scenario-based approach to test six propositions in which habitat is lost preferentially based on patch characteristics. We use eight bird and two amphibian species as indicator species. To compare scenarios, we present a method combining the output from a metapopulation model with measures of social impacts of land-cover change in a multiple criteria decision analysis. We also determine whether a habitat loss threshold exists, below which small loss of habitat can lead to large loss of species’ occupancy.

Results

We found that, of the scenarios presented, preferentially losing common habitats and smaller patches was least detrimental for both ecological and social factors. Threshold effects were found for all but the generalist bird species.

Conclusions

We have outlined a workflow which allows for transparent, repeatable comparison between landscapes. This workflow can be used to compare urban landscape plans, or to develop general understanding of the impacts of different forms of habitat loss. Reassuringly, the recommendations based on the scenarios presented are in keeping with received conservation wisdom: to prioritise larger and/or rarer patches.
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18.

Context

Habitat loss, fragmentation and degradation are widespread drivers of biodiversity decline. Understanding how habitat quality interacts with landscape context, and how they jointly affect species in human-modified landscapes, is of great importance for informing conservation and management.

Objectives

We used a whole-ecosystem manipulation experiment in the Brazilian Amazon to investigate the relative roles of local and landscape attributes in affecting bat assemblages at an interior-edge-matrix disturbance gradient.

Methods

We surveyed bats in 39 sites, comprising continuous forest (CF), fragments, forest edges and intervening secondary regrowth. For each site, we assessed vegetation structure (local-scale variable) and, for five focal scales, quantified habitat amount and four landscape configuration metrics.

Results

Smaller fragments, edges and regrowth sites had fewer species and higher levels of dominance than CF. Regardless of the landscape scale analysed, species richness and evenness were mostly related to the amount of forest cover. Vegetation structure and configurational metrics were important predictors of abundance, whereby the magnitude and direction of response to configurational metrics were scale-dependent. Responses were ensemble-specific with local-scale vegetation structure being more important for frugivorous than for gleaning animalivorous bats.

Conclusions

Our study indicates that scale-sensitive measures of landscape structure are needed for a more comprehensive understanding of the effects of fragmentation on tropical biota. Although forest fragments and regrowth habitats can be of conservation significance for tropical bats our results further emphasize that primary forest is of irreplaceable value, underlining that their conservation can only be achieved by the preservation of large expanses of pristine habitat.
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19.

Context

Species site-occupancy patterns may be influenced by habitat variables at both local and landscape scales. Although local habitat variables influence whether the site is suitable for a given species, the broader landscape context can also influence site occupancy, particularly for species that are sensitive to land-use change.

Objectives

To examine the relative importance of local versus landscape variables in explaining site occupancy of eight bat species within the Brazilian Cerrado, a Neotropical savanna that is experiencing widespread habitat loss and fragmentation.

Methods

Bats were surveyed within 16 forest patches over two years. We used a multi-model information-theoretic approach, adjusted for species detection bias, to assess whether landscape variables (percent cover and number of patches of natural vegetation within a 2- and 8-km radius of each forest site) or local site variables (canopy cover, understory height, number of trees, and number of lianas) best explained site occupancy in each species.

Results

Landscape variables were among the best models (ΔAICc or ΔQAICc < 2) for four species (top-ranked model for black myotis), whereas local variables were among the best for five species (top-ranked model for vampire bats). Neither local nor landscape variables explained site occupancy in two frugivorous species.

Conclusion

Species associated with a particular habitat type will not respond similarly to the amount, distribution or relative suitability of that habitat, or even at the same scale. This reinforces the challenge of species distribution modelling, especially in the context of forecasting species’ responses to future land-use or climate-change scenarios.
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20.

Context

Theory predicts that habitat loss and fragmentation may have drastic consequences on species’ interactions. To date, however, little empirical evidence exists on the strength of interspecific competition in shaping animal communities in fragmented landscapes.

Objectives

Our aim was to measure the degree of ongoing competitive interference between species in fragmented landscapes. Our model system was the community of ground-dwelling rodents in deciduous woodlands in central Italy, composed of a habitat generalist species (Apodemus sylvaticus) and two forest specialists (Apodemus flavicollis and Myodes glareolus). Our objectives were to test whether species were segregated among forest patches and whether spatial segregation was determined by interspecific competition or habitat and resource availability.

Methods

We surveyed the populations inhabiting 29 woodland patches in a highly fragmented landscape using a capture-mark-recapture protocol, capturing >4500 individuals. First we modelled species’ distribution as a function of habitat, resource availability and landscape variables. The second stage of our analyses involved measuring the response of vital rate parameters (body mass, reproduction, survival, recruitment, population density) to competitor density.

Results

The relative distribution of species reflected a spatial segregation of habitat generalists and specialists according to habitat quality, cover and connectivity. Interspecific competition mainly affected individual level vital rates, whereas we found no substantial effects at the population level.

Conclusions

Competitive exclusion of specialist species by generalist species was occurring. However, when compared to other factors such as habitat connectivity and resource availability, interspecific competition played a relatively minor role in shaping the studied community.
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