Landscape size affects the relative importance of habitat amount,habitat fragmentation,and matrix quality on forest birds

It is important to understand the relative effects of landscape habitat loss, habitat fragmentation, and matrix quality on biodiversity, so that potential management options can be appropriately ranked. However, their effects and relative importance may change with the size of the landscape considered because the multiple (and potentially conflicting) ecological processes that are influenced by landscape structure occur at different spatial scales (e.g. dispersal, predation, foraging). We estimated the relative effects of habitat loss, habitat fragmentation, and matrix quality (measured as the amount of forest, the proportion of forest area contained in large core forests, and the density of roads respectively) on fragmentation‐sensitive forest birds in southern Ontario, Canada using a range of landscape sizes (0.8–310 km²). We used three complementary statistical approaches to estimate relative effects of these correlated landscape factors – 1) multiple regression, 2) information theoretic (AIC) estimates of the most parsimonious model, and 3) multi‐model inference to average effects across all supported models. We controlled for spatial autocorrelation, local habitat, roadside sampling bias, time of day, season, habitat heterogeneity, and the interaction between the effects of habitat amount and fragmentation. We found that relative effects of habitat amount and fragmentation were scale dependent; habitat amount had a consistently positive effect that was consistent over more than two orders of magnitude in landscape area (~1–300 km²). In contrast, the effects of habitat fragmentation depended on the size of the landscape considered. Indeed, for veery Catharus fuscescens, habitat fragmentation had positive effects at one scale and negative effects at another. The effects of matrix quality were generally weak and changed little with scale. For the number of fragmentation sensitive species and the presence of veery, habitat amount was most important in large landscapes and habitat fragmentation in small landscapes but for the presence of ovenbird Seiurus aurocapilla, habitat amount was most important at all scales.     

生境破碎化对动物种群存活的影响

生境破碎是生物多样性下降的主要原因之一。通常以岛屿生物地理学、异质种群生物学和景观生态学的理论来解释不同空间尺度中生境破碎化的生态学效应。生境破碎化引起面积效应、隔离效应和边缘效应。这些效应通过影响动物种群的绝灭阈值、分布和多度、种间关系以及生态系统过程,最终影响动物种群的存活。野外研究表明,破碎化对动物的影响,因物种、生境类型和地理区域不同而有所变化,因此,预测物种在破碎生境中的存活比较困难。研究热点集中于：确定生境面积损失和生境斑块的空间格局对破碎景观中物种绝灭的相对影响,破碎景观中物种的适宜生境比例和绝灭阈值,异质种群动态以及生态系统的生态过程。随着3S技术的发展,生境破碎化模型趋于复杂,而发展有效的模型和验证模型将成为一项富有挑战性的任务。     

The Relationship between Habitat Loss and Fragmentation during Urbanization: An Empirical Evaluation from 16 World Cities

Urbanization results in habitat loss and habitat fragmentation concurrently, both influencing biodiversity and ecological processes. To evaluate these impacts, it is important to understand the relationships between habitat loss and habitat fragmentation per se (HLHF) during urbanization. The objectives of this study were two-fold: 1) to quantify the different forms of the HLHF relationship during urbanization using multiple landscape metrics, and 2) to test the validity of the HLHF relations reported in the literature. Our analysis was based on a long-term urbanization dataset (1800–2000) of 16 large cities from around the world. Habitat area was represented as the percentage of non-built-up area in the landscape, while habitat fragmentation was measured using several landscape metrics. Our results show that the relationship between habitat loss and habitat fragmentation during urbanization is commonly monotonic—linear, exponential, or logarithmic, indicating that the degree of habitat fragmentation per se increases with habitat loss in general. We compared our results with 14 hypothesized HLHF relationships based on simulated landscapes found in the literature, and found that four of them were consistent with those of urbanization, whereas the other ten were not. Also, we identified six new HLHF relationships when fragmentation was measured by total core area, normalized total core area, patch density, edge density and landscape shape index, respectively. In addition, our study demonstrated that the “space-for-time” approach, frequently used in ecology and geography, generated specious HLHF relationships, suggesting that this approach is largely inappropriate for analyses of urban landscapes that are highly heterogeneous in space and unusually contingent in dynamics. Our results show both generalities and idiosyncrasies of the HLHF relationship, providing new insights for assessing ecological effects of urbanization.     

Rethinking the conceptual foundations of habitat fragmentation research

The conceptual foundations of habitat fragmentation research have not kept pace with empirical advances in our understanding of species responses to landscape change, nor with theoretical advances in the wider disciplines of ecology. There is now real debate whether explicit recognition of ‘habitat fragmentation’ as an over‐arching conceptual domain will stimulate or hinder further progress toward understanding and mitigating the effects of landscape change. In this paper, we critically challenge the conceptual foundations of the discipline, and attempt to derive an integrated perspective on the best way to advance mechanistic understanding of fragmentation processes. We depict the inherent assumptions underlying the discipline as a ‘conceptual phase space’ of contrasting false dichotomies in fragmentation ‘problem space’. In our opinion, the key determinant of whether ‘habitat fragmentation’ can remain a cohesive framework lies in the concept of ‘interdependence’: 1) interdependence of landscape effects on species and 2) interdependence of species responses to landscape change. If there is non‐trivial interdependence among the various sub‐components of habitat fragmentation, or non‐trivial interdependence among species responses to landscape change, then there will be real heuristic value in ‘habitat fragmentation’ as a single conceptual domain. At present, the current paradigms entrenched in the fragmentation literature are implicitly founded on strict independence of landscape effects (e.g. the debate about the independent effects of habitat loss versus fragmentation per se) and strict independence of species responses (e.g. the individualistic species response models underpinning landscape continuum models), despite compelling evidence for interdependence in both effects and responses to fragmentation. We discuss how strong ‘interdependence’ of effects and responses challenges us to rethink long‐held views, and re‐cast the conceptual foundations of habitat fragmentation in terms of spatial context‐dependence in the effects of multiple interacting spatial components of fragmentation, and community context‐dependence in the responses of multiple interacting species to landscape change.     

Characterising extinction debt following habitat fragmentation using neutral theory

Habitat loss leads to species extinctions, both immediately and over the long term as ‘extinction debt’ is repaid. The same quantity of habitat can be lost in different spatial patterns with varying habitat fragmentation. How this translates to species loss remains an open problem requiring an understanding of the interplay between community dynamics and habitat structure across temporal and spatial scales. Here we develop formulas that characterise extinction debt in a spatial neutral model after habitat loss and fragmentation. Central to our formulas are two new metrics, which depend on properties of the taxa and landscape: ‘effective area’, measuring the remaining number of individuals and ‘effective connectivity’, measuring individuals’ ability to disperse through fragmented habitat. This formalises the conventional wisdom that habitat area and habitat connectivity are the two critical requirements for long‐term preservation of biodiversity. Our approach suggests that mechanistic fragmentation metrics help resolve debates about fragmentation and species loss.     

Direct versus indirect effects of habitat fragmentation on community patterns in experimental landscapes

Habitat area and fragmentation are confounded in many ecological studies investigating fragmentation effects. We thus devised an innovative experiment founded on fractal neutral landscape models to disentangle the relative effects of habitat area and fragmentation on arthropod community patterns in red clover (Trifolium pratense). The conventional approach in experimental fragmentation studies is to adjust patch size and isolation to create different landscape patterns. We instead use fractal distributions to adjust the overall amount and fragmentation of habitat independently at the scale of the entire landscape, producing different patch properties. Although habitat area ultimately had a greater effect on arthropod abundance and diversity in this system, we found that fragmentation had a significant effect in clover landscapes with ≤40 % habitat. Landscapes at these lower habitat levels were dominated by edge cells, which had fewer arthropods and lower richness than interior cells. Fragmentation per se did not have a direct effect on local-scale diversity, however, as demonstrated by the lack of a broader landscape effect (in terms of total habitat area and fragmentation) on arthropods within habitat cells. Fragmentation—through the creation of edge habitat—thus had a strong indirect effect on morphospecies richness and abundance at the local scale. Although it has been suggested that fragmentation should be important at low habitat levels (≤20–30 %), we show that fragmentation per se is significant only at intermediate (40 %) levels of habitat, where edge effects were neither too great (as at lower levels of habitat) nor too weak (as at higher levels of habitat).     

Avifaunal responses to habitat fragmentation in the threatened littoral forests of south-eastern Madagascar

Aim Madagascar's lowland forests are both rich in endemic taxa and considered to be seriously threatened by deforestation and habitat fragmentation. However, very little is known about how these processes affect biodiversity on the island. Herein, we examine how forest bird communities and functional groups have been affected by fragmentation at both patch and landscape scales, by determining relationships between species richness and individual species abundance and patch and landscape mosaic metrics. Location Littoral forest remnants within south‐eastern Madagascar. Methods We sampled 30 littoral forest remnants in south‐eastern Madagascar, within a landscape mosaic dominated by Erica spp. heathland. We quantified bird community composition within remnants of differing size, shape and isolation, by conducting point counts in November–December in 2001 and October–November 2002. Each remnant was characterized by measures of remnant area, remnant shape, isolation, and surrounding landscape complexity. We used step‐wise regression to test the relationship between bird species richness and landscape structural elements, after correcting for sampling effort. Relationships between bird species abundances and the landscape variables were investigated with Canonical Correspondence Analysis and binomial logistic regression modelling. Results Bird species richness and forest‐dependent bird species richness were significantly (P < 0.01) explained by remnant area but not by any measure of isolation or landscape complexity. The majority of forest‐dependent species had significant relationships with remnant area. Minimum area requirements for area‐sensitive species ranged from 15 to 150 ha, with the majority of species having area requirements > 30 ha. Surprisingly, there was no relationship between bird body size and minimum area requirement. Forest‐dependent canopy insectivorous species and large canopy frugivorous species were the most sensitive functional groups, with > 90% species sensitivity within each group. The distribution of four forest‐dependent species also appeared to be related to remnant shape where remnant area was < 100 ha. Main conclusions The majority of forest‐dependent species, including many that are considered widespread and common, were found to have significant relationships with fragment size, indicating that they are sensitive to processes associated with habitat loss and fragmentation. As deforestation and habitat fragmentation remain serious problems on the island, it follows that many forest‐dependent bird species will decline in abundance and become locally extinct. At the regional scale, we urge that large (> 200 ha) blocks of littoral forest are awarded protected status to preserve their unique bird community.     

Landscape structure effects on forest plant diversity at local scale: Exploring the role of spatial extent

Since landscape attributes show different patterns at different spatial extents, it is fundamental to identify how the relation between landscape structure and plant species diversity at local scale varies with scale. Then, it is fundamental to assess the appropriate extent at which landscape factors affect plant species richness at the local scale. To investigate this relation, data on plant species richness of forest communities at plot scale were extracted from a large data set and landscape metrics were calculated around the same plots for a range of extents (250–3000 m). Then, multiple regression models and variance partitioning techniques were applied to assess the amount of variance explained by the landscape metrics on plant species richness for a range of extents. In general, we found that increasing extent of the surrounding landscape analyzed, improved the strength of relationship between the landscape metrics and the properties of plant communities at plot scale. The medium-large extent was most informative as it combined a decent total variance explained with high variance explained by the pure fractions of complexity, fragmentation and disturbance and the minimum of collinearity. In conclusion, we found that it is possible and beneficial to identify a specific extent, where the redundancy in the predictor variables is minimized and the explanatory power of the pure fractions (or single groups) maximized, when examining landscape structure effects on local plant species richness.     

Landscape-level comparison of genetic diversity and differentiation in a small mammal inhabiting different fragmented landscapes of the Brazilian Atlantic Forest

Habitat loss and fragmentation can have detrimental effects on all levels of biodiversity, including genetic variation. Most studies that investigate genetic effects of habitat loss and fragmentation focus on analysing genetic data from a single landscape. However, our understanding of habitat loss effects on landscape-wide patterns of biodiversity would benefit from studies that are based on quantitative comparisons among multiple study landscapes. Here, we use such a landscape-level study design to compare genetic variation in the forest-specialist marsupial Marmosops incanus from four 10,000-hectare Atlantic forest landscapes which differ in the amount of their remaining native forest cover (86, 49, 31, 11 %). Additionally, we used a model selection framework to evaluate the influence of patch characteristics on genetic variation within each landscape. We genotyped 529 individuals with 12 microsatellites to statistically compare estimates of genetic diversity and genetic differentiation in populations inhabiting different forest patches within the landscapes. Our study indicates that before the extinction of the specialist species (here in the 11 % landscape) genetic diversity is significantly reduced in the 31 % landscape, while genetic differentiation is significantly higher in the 49 and 31 % landscapes compared to the 86 % landscape. Results further provide evidence for non-proportional responses of genetic diversity and differentiation to increasing habitat loss, and suggest that local patch isolation impacts gene flow and genetic connectivity only in the 31 % landscape. These results have high relevance for analysing landscape genetic relationships and emphasize the importance of landscape-level study designs for understanding habitat loss effects on all levels of biodiversity.     

Effects of habitat loss, habitat fragmentation, and isolation on the density, species richness, and distribution of ladybeetles in manipulated alfalfa landscapes

Abstract.  1. Habitat loss and fragmentation are the main causes of changes in the distribution and abundance of organisms, and are usually considered to negatively affect the abundance and species richness of organisms in a landscape. Nevertheless, habitat loss and fragmentation have often been confused, and the reported negative effects may only be the result of habitat loss alone, with habitat fragmentation having nil or even positive effects on abundance and species richness.
2. Manipulated alfalfa micro-landscapes and coccinellids (Coleoptera: Coccinellidae) are used to test the effects habitat loss (0% or 84%), fragmentation (4 or 16 fragments), and isolation (2 or 6 m between fragments) on the density, species richness, and distribution of native and exotic species of coccinellids.
3. Generally, when considering only the individuals in the remaining fragments, habitat loss had variable effects while habitat fragmentation had a positive effect on the density of two species of coccinellids and on species richness, but did not affect two other species. Isolation usually had no effect. When individuals in the whole landscape were considered, negative effects of habitat loss became apparent for most species, but the positive effects of fragmentation remained only for one species.
4. Native and exotic species of coccinellids did not segregate in the different landscapes, and strong positive associations were found most often in landscapes with higher fragmentation and isolation.
5. The opposing effects of habitat loss and fragmentation may result in a nil global effect; therefore it is important to separate their effects when studying populations in fragmented landscapes.     

Landscape genetics and limiting factors

Population connectivity is mediated by the movement of organisms or propagules through landscapes. However, little is known about how variation in the pattern of landscape mosaics affects the detectability of landscape genetic relationships. The goal of this paper is to explore the impacts of limiting factors on landscape genetic processes using simulation modeling. We used spatially explicit, individual-based simulation modeling to quantify the effects of habitat area, fragmentation and the contrast in resistance between habitat and non-habitat on the apparent strength and statistical detectability of landscape genetic relationships. We found that landscape genetic effects are often not detectable when habitat is highly connected. In such situations landscape structure does not limit gene flow. We also found that contrast in resistance values between habitat and non-habitat interacts with habitat extensiveness and fragmentation to affect detectability of landscape genetic relationships. Thus, the influence of landscape features critical to providing connectivity may not be detectable if gene flow is not limited by spatial patterns or resistance contrast of these features. We developed regression equations that reliably predict whether or not isolation by resistance will be detected independently of isolation by distance as a function of habitat fragmentation and contrast in resistance between habitat and non-habitat.     

Reversing habitat loss: deciduous habitat fragmentation matters to birds in a larch plantation matrix

Whereas matrix management has recently been suggested as a useful tool in biodiversity conservation, patterns and processes within matrices remain unknown. We examined the effects of the loss and fragmentation (configuration) of original deciduous habitats on birds in larch plantation matrix in the winter and during the breeding season in a montane region in Nagano Prefecture, central Japan. Birds were counted using a plot-count method from 32 (winter) and 46 (breeding) matrix (plantation) sites with a range of surrounding habitat loss and fragmentation at a 1600-m scale. Birds were assigned to species groups based on ecological traits, and three groups for which larch plantation would function as low-quality matrix were analyzed. Bird occurrences were explained primarily by habitat structure. Although the effects of landscape structure were less than those of habitat structure, three of five groups across the two seasons experienced significant landscape effects. All effects were of habitat fragmentation (scatteredness of deciduous habitats), i.e. two groups (flycatchers in the breeding season and tree nesters in both seasons) frequently occurred in matrix surrounded by highly scattered deciduous habitats. However, most of these effects were confounded by local habitat structure. That is, the variation in bird occurrences explained purely by fragmentation variables was <6%. Nonetheless, because these effects were marginally significant at p<0.10, there was some support for fragmentation effects after covariation with habitat structure was considered. Based on these results and associated theoretical studies, we hypothesized that habitat fragmentation leads to loss of individuals and is more important than habitat loss in landscapes with a structurally similar matrix. We also hypothesized that matrix within landscapes with highly scattered habitats should be managed with high priority.     

Responses of 'resistant' vertebrates to habitat loss and fragmentation: the importance of niche breadth and range boundaries

Abstract. An ability to predict species' sensitivities to habitat loss and fragmentation has important conservation implications, and numerous hypotheses have been proposed to explain interspecific differences observed in human-dominated landscapes. We used occupancy data collected on 32 species of vertebrates (16 mammals and 16 amphibians) in an agricultural landscape of Indiana, USA, to compare hypotheses that focus on different causal mechanisms underlying interspecific variation in responses to habitat alteration: (1) body size; (2) morphology and development; (3) behaviour; (4) niche breadth; (5) proximity to range boundary; and multiple-process models combining main effects and interactions of hypotheses (1)–(2) and (4)–(5). The majority of habitat alteration occurred over a century ago and coincided with extinction of several species; thus, our study dealt only with variation in responses of extant species that often are considered 'resistant' to human modifications of native habitat. Corrected Akaike scores and Akaike weights provided strongest support for models incorporating niche breadth and proximity to range boundary. Measures of dietary and habitat breadth obtained from the literature were negatively correlated with sensitivity to habitat alteration. Additionally, greater sensitivity was observed for species occurring at the periphery of their geographical ranges, especially at northern or western margins. Body size, morphological, developmental and behavioural traits were inferior predictors of tolerance to fragmentation for the species and landscape we examined. Our findings reinforce the importance of niche breadth as a predictor of species' responses to habitat alteration. They also highlight the importance of viewing the effects of habitat loss and fragmentation in a landscape within a biogeographical context that considers a species' level of adaptation to local environmental conditions.     

Tolerance to habitat fragmentation influences the colonization of new habitat by forest birds

We examined the relationship between the ability of bird species to persist in fragmented forests and their ability to colonize new forest habitat. Using a long-term data set on the colonization of a forest plantation, we tested the hypothesis that bird species tolerant to habitat fragmentation would detect and colonize the new habitat faster than intolerant species. The forest plantation under study is situated on an area of land reclaimed from the sea (a polder) in the central part of The Netherlands. We constructed an index of tolerance to habitat fragmentation and included it as a predictor variable in a set of three logistic regression models that compared the probability of colonization over four consecutive time periods. After controlling statistically for the effects of regional incidence, preferred habitat and life-history characteristics, there was a significant effect of tolerance to fragmentation on the ability of species to colonize the plantation, and a marginal effect on the timing of colonization. We then examined the effect of the same index of tolerance to fragmentation on colonization patterns over a larger spatial scale. Multivariate regression models showed that the proportion of three polders of different ages occupied by forest bird species was dependent upon the regional incidence of a species, its preferred habitat and its tolerance to fragmentation. The results support the hypothesis that species tolerant to habitat fragmentation detect and colonize new habitat faster than those intolerant to habitat fragmentation.     

A species‐centered approach for uncovering generalities in organism responses to habitat loss and fragmentation

Theoretical models predict strong influences of habitat loss and fragmentation on species distributions and demography, but empirical studies have shown relatively inconsistent support across species and systems. We argue that species’ responses to landscape‐scale habitat loss and fragmentation are likely to appear less idiosyncratic if it is recognized that species perceive the same landscapes in different ways. We present a new quantitative approach that uses species distribution models (SDMs) to measure landscapes (e.g. patch size, isolation, matrix amount) from the perspective of individual species. First, we briefly summarize the few efforts to date demonstrating that once differences in habitat distributions are controlled, consistencies in species’ responses to landscape structure emerge. Second, we present a detailed example providing step‐by‐step methods for application of a species‐centered approach using freely available land‐cover data and recent statistical modeling approaches. Third, we discuss pitfalls in current applications of the approach and recommend avenues for future developments. We conclude that the species‐centered approach offers considerable promise as a means to test whether sensitivity to habitat loss and fragmentation is mediated by phylogenetic, ecological, and life‐history traits. Cross‐species generalities in responses to habitat loss and fragmentation will be challenging to uncover unless landscape mosaics are defined using models that reflect differing species‐specific distributions, functional connectivity, and domains of scale. The emergence of such generalities would not only enhance scientific understanding of biotic processes driving fragmentation effects, but would allow managers to estimate species sensitivities in new regions.     

Direct and indirect effects of habitat area and matrix composition on species interactions among flower-visiting insects

Traditionally, ecologists have focused on direct effects of habitat area and arrangement on individual species or pairwise species interactions. Indirect effects of habitat heterogeneity on multiple interacting species are often neglected or lack experimental support. In a factorial field experiment, we explored the direct and indirect effects of habitat area, fragmentation, and matrix composition on a community of flower-visiting insects in red clover. Habitat area, fragmentation, and density of inflorescences of red clover all influenced the insect pollinators and, in turn, the production of clover seeds. The strongest direct effect was on pollinator visitation, which was substantially higher in small clover patches surrounded by bare-ground than in larger patches embedded within grass. Structural equation modeling indicated that the observed matrix-dependent changes in pollinator visitation propagated across a tri-trophic system: higher visitation rates positively correlated with a higher seed set, which in turn was positively correlated with abundances of seed predators and their parasitoids. Therefore, this study suggests that habitat area and matrix composition can strongly influence the structure of species interaction webs through indirect effects, and also emphasizes that these effects can be propagated through mutualistic as well as trophic interactions.     

Independent effects of habitat loss,habitat fragmentation and structural connectivity on forest‐dependent birds

Aim Habitat loss and fragmentation are amongst the greatest threats to biodiversity world‐wide. However, there is still little evidence on the relative influence of these two distinct processes on biodiversity, and no study, to date, has investigated the independent contribution of structural connectivity in addition to habitat loss and fragmentation. The aim of this study is to evaluate the independent effects of habitat loss (the decrease in total amount of habitat), habitat fragmentation per se (habitat subdivision) and structural connectivity (in the form of hedgerow networks) on the distribution of seven resident forest‐dependent birds in central Italy. Location Central Italy. Methods We strategically selected 30 landscapes (each of 16 km² in size) with decreasing total amount of forest cover and with contrasting configuration of patches and contrasting lengths of hedgerow networks. Presence/absence of birds in each landscape unit was studied through point counts. Results The amount of forest cover in the landscape had the strongest relative influence on birds’ occupancy, whilst habitat subdivision played a negligible role. Structural connectivity and the geographic position of the landscape unit played a relatively important role for four species. Main conclusions Our study shows the importance of disentangling the contribution of different landscape properties in determining distribution patterns. Our results are consistent with the fact that halting habitat loss and carrying out habitat restoration should be conservation priorities, since habitat loss is the main factor affecting the distribution of the target species; implementation of structural connectivity through hedgerows, instead, should be evaluated with caution since its contribution is secondary to the predominant role of habitat loss.     

Spatial heterogeneity and habitat configuration overcome habitat composition influences on alpha and beta mammal diversity

The effects of habitat fragmentation on different taxa and ecosystems are subject to intense debate, and disentangling them is of utmost importance to support conservation and management strategies. We evaluated the importance of landscape composition and configuration, and spatial heterogeneity to explain α- and β-diversity of mammals across a gradient of percent woody cover and land use diversity. We expected species richness to be positively related to all predictive variables, with the strongest relationship with landscape composition and configuration, and spatial heterogeneity respectively. We also expected landscape to influence β-diversity in the same order of importance expected for species richness, with a stronger influence on nestedness due to deterministic loss of species more sensitive to habitat disturbance. We analyzed landscape structure using: (a) landscape metrics based on thematic maps and (b) image texture of a vegetation index. We compared a set of univariate explanatory models of species richness using AIC, and evaluated how dissimilarities in landscape composition and configuration and spatial heterogeneity affect β-diversity components using a Multiple Regression on distance Matrix. Contrary with our expectations, landscape configuration was the main driver of species richness, followed by spatial heterogeneity and last by landscape composition. Nestedness was explained, in order of importance, by spatial heterogeneity, landscape configuration, and landscape composition. Although conservation policies tend to focus mainly on habitat amount, we advocate that landscape management must include strategies to preserve and improve habitat quality and complexity in natural patches and the surrounding matrix, enabling landscapes to harbor high species diversity.     

Critical thresholds associated with habitat loss: a review of the concepts, evidence, and applications

A major conservation concern is whether population size and other ecological variables change linearly with habitat loss, or whether they suddenly decline more rapidly below a “critical threshold” level of habitat. The most commonly discussed explanation for critical threshold responses to habitat loss focus on habitat configuration. As habitat loss progresses, the remaining habitat is increasingly fragmented or the fragments are increasingly isolated, which may compound the effects of habitat loss. In this review we also explore other possible explanations for apparently nonlinear relationships between habitat loss and ecological responses, including Allee effects and time lags, and point out that some ecological variables will inherently respond nonlinearly to habitat loss even in the absence of compounding factors. In the literature, both linear and nonlinear ecological responses to habitat loss are evident among simulation and empirical studies, although the presence and value of critical thresholds is influenced by characteristics of the species (e.g. dispersal, reproduction, area/edge sensitivity) and landscape (e.g. fragmentation, matrix quality, rate of change). With enough empirical support, such trends could be useful for making important predictions about species' responses to habitat loss, to guide future research on the underlying causes of critical thresholds, and to make better informed management decisions. Some have seen critical thresholds as a means of identifying conservation targets for habitat retention. We argue that in many cases this may be misguided, and that the meaning (and utility) of a critical threshold must be interpreted carefully and in relation to the response variable and management goal. Despite recent interest in critical threshold responses to habitat loss, most studies have not used any formal statistical methods to identify their presence or value. Methods that have been used include model comparisons using Akaike information criterion (AIC) or t‐tests, and significance testing for changes in slope or for polynomial effects. The judicious use of statistics to help determine the shape of ecological relationships would permit greater objectivity and more comparability among studies.     

Extinction–immigration dynamics lag behind environmental filtering in shaping the composition of tropical dry forests within a changing landscape

The impact of rapid habitat loss and fragmentation on biodiversity is a major issue. However, we still lack an integrative understanding of how these changes influence biodiversity dynamics over time. In this study, we investigate the effects of these changes in terms of both niche-based and neutral dynamics. We hypothesize that habitat loss has delayed effects on neutral immigration–extinction dynamics, while edge effects and environmental heterogeneity in habitat patches have rapid effects on niche-based dynamics. We analyzed taxonomic and functional composition of 100 tree communities in a tropical dry forest landscape of New-Caledonia subject to habitat loss and fragmentation. We designed an original, process-based simulation framework, and performed Approximate Bayesian Computation to infer the influence of niche-based and neutral processes. Then, we performed partial regressions to evaluate the relationships between inferred parameter values of communities and landscape metrics (distance to edge, patch area, and habitat amount around communities), derived from either recent or past (65 yr ago) aerial photographs, while controlling for the effect of soil and topography. We found that landscape structure influences both environmental filtering and immigration. Immigration rate was positively related to past habitat amount surrounding communities. In contrast, environmental filtering was mostly affected by present landscape structure and mainly influenced by edge vicinity and topography. Our results highlight that landscape changes have contrasting spatio-temporal influences on niche-based and neutral assembly dynamics. First, landscape-level habitat loss and community isolation reduce immigration and increase demographic stochasticity, resulting in slow decline of local species diversity and extinction debt. Second, recent edge creation affects environmental filtering, incurring rapid changes in community composition by favoring species with edge-adapted strategies. Our study brings new insights about temporal impacts of landscape changes on biodiversity dynamics. We stress that landscape history critically influences these dynamics and should be taken into account in conservation policies.