Anthropogenic landscape change promotes asymmetric dispersal and limits regional patch occupancy in a spatially structured bird population

1. Local extinctions in habitat patches and asymmetric dispersal between patches are key processes structuring animal populations in heterogeneous environments. Effective landscape conservation requires an understanding of how habitat loss and fragmentation influence demographic processes within populations and movement between populations. 2. We used patch occupancy surveys and molecular data for a rainforest bird, the logrunner (Orthonyx temminckii), to determine (i) the effects of landscape change and patch structure on local extinction; (ii) the asymmetry of emigration and immigration rates; (iii) the relative influence of local and between-population landscapes on asymmetric emigration and immigration; and (iv) the relative contributions of habitat loss and habitat fragmentation to asymmetric emigration and immigration. 3. Whether or not a patch was occupied by logrunners was primarily determined by the isolation of that patch. After controlling for patch isolation, patch occupancy declined in landscapes experiencing high levels of rainforest loss over the last 100 years. Habitat loss and fragmentation over the last century was more important than the current pattern of patch isolation alone, which suggested that immigration from neighbouring patches was unable to prevent local extinction in highly modified landscapes. 4. We discovered that dispersal between logrunner populations is highly asymmetric. Emigration rates were 39% lower when local landscapes were fragmented, but emigration was not limited by the structure of the between-population landscapes. In contrast, immigration was 37% greater when local landscapes were fragmented and was lower when the between-population landscapes were fragmented. Rainforest fragmentation influenced asymmetric dispersal to a greater extent than did rainforest loss, and a 60% reduction in mean patch area was capable of switching a population from being a net exporter to a net importer of dispersing logrunners. 5. The synergistic effects of landscape change on species occurrence and asymmetric dispersal have important implications for conservation. Conservation measures that maintain large patch sizes in the landscape may promote asymmetric dispersal from intact to fragmented landscapes and allow rainforest bird populations to persist in fragmented and degraded landscapes. These sink populations could form the kernel of source populations given sufficient habitat restoration. However, the success of this rescue effect will depend on the quality of the between-population landscapes.     

Mammal responses to matrix development intensity

The landscape matrix is increasingly being recognized as important to biodiversity conservation. The nature of the matrix impacts the persistence of species in human‐modified landscapes through its pervasive influence on adjacent habitat and through the habitat value of the matrix itself. However, previous studies have not isolated the effects of the matrix from the effects of other aspects of landscape modification, such as habitat loss and fragmentation, and much remains to be understood about the independent impact of the matrix on wildlife. We investigated the effects of the matrix on mammal abundance and landscape use in south‐east Queensland, Australia. Mammals were surveyed in patch ‘core’, patch ‘edge’ and ‘matrix’ landscape elements along a rural–suburban gradient of matrix development intensity quantified by a weighted road‐length metric, which was significantly correlated with housing density, while controlling for potentially confounding patch and landscape attributes. Response to increasing matrix development intensity was highly species‐specific. Several native species declined in abundance; however, others were more resilient to moderate levels of matrix intensity, one species increased in abundance, and at least one species appeared unaffected by matrix intensity. Native species richness peaked at moderate levels of matrix development intensity. Exotic species richness and feral predators increased with matrix intensity and were negatively correlated with native species. Species response to matrix intensity appeared related to their use of edge or matrix habitat. An ability to use the matrix per se, however, may not translate into an ability to persist in a landscape where development substantially reduces the habitat or movement value of the matrix.     

Landscape modification and habitat fragmentation: a synthesis

Landscape modification and habitat fragmentation are key drivers of global species loss. Their effects may be understood by focusing on: (1) individual species and the processes threatening them, and (2) human-perceived landscape patterns and their correlation with species and assemblages. Individual species may decline as a result of interacting exogenous and endogenous threats, including habitat loss, habitat degradation, habitat isolation, changes in the biology, behaviour, and interactions of species, as well as additional, stochastic threats. Human-perceived landscape patterns that are frequently correlated with species assemblages include the amount and structure of native vegetation, the prevalence of anthropogenic edges, the degree of landscape connectivity, and the structure and heterogeneity of modified areas. Extinction cascades are particularly likely to occur in landscapes with low native vegetation cover, low landscape connectivity, degraded native vegetation and intensive land use in modified areas, especially if keystone species or entire functional groups of species are lost. This review (1) demonstrates that species-oriented and pattern-oriented approaches to understanding the ecology of modified landscapes are highly complementary, (2) clarifies the links between a wide range of interconnected themes, and (3) provides clear and consistent terminology. Tangible research and management priorities are outlined that are likely to benefit the conservation of native species in modified landscapes around the world.     

The effects of landscape fragmentation on pollination dynamics: absence of evidence not evidence of absence

Animal‐mediated pollination is essential for both ecosystem services and conservation of global biodiversity, but a growing body of work reveals that it is negatively affected by anthropogenic disturbance. Landscape‐scale disturbance results in two often inter‐related processes: (1) habitat loss, (2) disruptions of habitat configuration (i.e. fragmentation). Understanding the relative effects of such processes is critical in designing effective management strategies to limit pollination and pollinator decline. We reviewed existing published work from 1989 to 2009 and found that only six of 303 studies considering the influence of landscape context on pollination separated the effects of habitat loss from fragmentation. We provide a synthesis of the current landscape, behavioural, and pollination ecology literature in order to present preliminary multiple working hypotheses explaining how these two landscape processes might independently influence pollination dynamics. Landscape disturbance primarily influences three components of pollination interactions: pollinator density, movement, and plant demography. We argue that effects of habitat loss on each of these components are likely to differ substantially from the effects of fragmentation, which is likely to be more complex and may influence each pollination component in contrasting ways. The interdependency between plants and animals inherent to pollination systems also has the possibility to drive cumulative effects of fragmentation, initiating negative feedback loops between animals and the plants they pollinate. Alternatively, due to their asymmetrical structure, pollination networks may be relatively robust to fragmentation. Despite the potential importance of independent effects of habitat fragmentation, its effects on pollination remain largely untested. We postulate that variation across studies in the effects of ‘fragmentation’ owes much to artifacts of the sampling regimes adopted, particularly (1) incorrectly separating fragmentation from habitat loss, and (2) mis‐matches in spatial scale between landscapes studied and the ecological processes of interest. The field of landscape pollination ecology could be greatly advanced through the consideration and quantification of the matrix, landscape functional connectivity, and pollinator movement behaviour in response to these elements. Studies designed to disentangle the independent effects of habitat loss and fragmentation are essential for gaining insight into landscape‐mediated pollination declines, implementing effective conservation measures, and optimizing ecosystem services in complex landscapes.     

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.     

The behavioural ecology and population dynamics of a cryptic ground‐dwelling mammal in an urban Australian landscape

Urbanization results in widespread habitat loss and fragmentation and generally has a negative impact upon native wildlife, in particular ground‐dwelling mammals. The northern brown bandicoot (Isoodon macrourus; Marsupialia: Peramelidae) is one of relatively few native Australian ground‐dwelling mammals that is able to survive within urbanized landscapes. As a consequence of extensive clearing and urban development within the city of Brisbane, bandicoots are now restricted to the mostly small (<10 ha) bushland fragments scattered across the city landscape. Our study examined the behavioural ecology of northern brown bandicoots within habitat fragments located on a major creek‐line, using mark‐recapture population monitoring and radio telemetry. Bandicoots at monitored sites were found to occur at high densities (typically one individual ha^?1), although one‐third of the populations were transient. Radio tracking revealed that bandicoots had relatively small home ranges (mean 1.5 ± 0.2 ha) comprised largely of bushland/grassland with dense, often weed‐infested ground cover. Bandicoots sheltered by day in these densely covered areas and also spent most time foraging there at night, although they occasionally ventured small distances to forage in adjacent maintained parklands and residential lawns. We suggest that introduced tall grasses and other weeds contribute to high habitat quality within riparian habitat fragments and facilitate the persistence of high density populations, comprised of individuals with small home ranges. The generalized dietary and habitat requirements of northern brown bandicoots, as well as a high reproductive output, undoubtedly facilitate the survival of the species in urban habitat fragments. Further research is required on other native mammal species in urbanized landscapes to gain a greater understanding of how best to conserve wildlife in these heavily modified environments.     

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.     

Landscape-level toxicant exposure mediates infection impacts on wildlife populations

Anthropogenic landscape modification such as urbanization can expose wildlife to toxicants, with profound behavioural and health effects. Toxicant exposure can alter the local transmission of wildlife diseases by reducing survival or altering immune defence. However, predicting the impacts of pathogens on wildlife across their ranges is complicated by heterogeneity in toxicant exposure across the landscape, especially if toxicants alter wildlife movement from toxicant-contaminated to uncontaminated habitats. We developed a mechanistic model to explore how toxicant effects on host health and movement propensity influence range-wide pathogen transmission, and zoonotic exposure risk, as an increasing fraction of the landscape is toxicant-contaminated. When toxicant-contaminated habitat is scarce on the landscape, costs to movement and survival from toxicant exposure can trap infected animals in contaminated habitat and reduce landscape-level transmission. Increasing the proportion of contaminated habitat causes host population declines from combined effects of toxicants and infection. The onset of host declines precedes an increase in the density of infected hosts in contaminated habitat and thus may serve as an early warning of increasing potential for zoonotic spillover in urbanizing landscapes. These results highlight how sublethal effects of toxicants can determine pathogen impacts on wildlife populations that may not manifest until landscape contamination is widespread.     

Urban landscapes can change virus gene flow and evolution in a fragmentation‐sensitive carnivore

Urban expansion has widespread impacts on wildlife species globally, including the transmission and emergence of infectious diseases. However, there is almost no information about how urban landscapes shape transmission dynamics in wildlife. Using an innovative phylodynamic approach combining host and pathogen molecular data with landscape characteristics and host traits, we untangle the complex factors that drive transmission networks of feline immunodeficiency virus (FIV) in bobcats (Lynx rufus). We found that the urban landscape played a significant role in shaping FIV transmission. Even though bobcats were often trapped within the urban matrix, FIV transmission events were more likely to occur in areas with more natural habitat elements. Urban fragmentation also resulted in lower rates of pathogen evolution, possibly owing to a narrower range of host genotypes in the fragmented area. Combined, our findings show that urban landscapes can have impacts on a pathogen and its evolution in a carnivore living in one of the most fragmented and urban systems in North America. The analytical approach used here can be broadly applied to other host–pathogen systems, including humans.     

Contrasting effects of mosaic structure on alpha and beta diversity of bird assemblages in a human‐modified landscape

Habitat loss and fragmentation are key processes causing biodiversity loss in human‐modified landscapes. Knowledge of these processes has largely been derived from measuring biodiversity at the scale of ‘within‐habitat’ fragments with the surrounding landscape considered as matrix. Yet, the loss of variation in species assemblages ‘among’ habitat fragments (landscape‐scale) may be as important a driver of biodiversity loss as the loss of diversity ‘within’ habitat fragments (local‐scale). We tested the hypothesis that heterogeneity in vegetation cover is important for maintaining alpha and beta diversity in human‐modified landscapes. We surveyed bird assemblages in eighty 300‐m‐long transects nested within twenty 1‐km² vegetation ‘mosaics’, with mosaics assigned to four categories defined by the cover extent and configuration of native eucalypt forest and exotic pine plantation. We examined bird assemblages at two spatial scales: 1) within and among transects, and 2) within and among mosaics. Alpha diversity was the mean species diversity within‐transects or within‐mosaics and beta diversity quantified the effective number of compositionally distinct transects or mosaics. We found that within‐transect alpha diversity was highest in vegetation mosaics defined by continuous eucalypt forest, lowest in mosaics of continuous pine plantation, and at intermediate levels in mosaics containing eucalypt patches in a pine matrix. We found that eucalypt mosaics had lower beta diversity than other mosaic types when ignoring relative abundances, but had similar or higher beta diversity when weighting with species abundances. Mosaics containing both pine and eucalypt forest differed in their bird compositional variation among transects, despite sharing a similar suite of species. This configuration effect at the mosaic scale reflected differences in vegetation composition among transects. Maintaining heterogeneity in vegetation cover could help to maintain variation among bird assemblages across landscapes, thus partially offsetting local‐scale diversity losses due to fragmentation. Critical to this is the retention of remnant native vegetation.     

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.     

Response of a small felid of conservation concern to habitat fragmentation

Habitat loss and fragmentation are major drivers of biodiversity loss. A key question, particularly relevant to carnivore conservation, is to which extent species are able to survive in human-modified landscapes. Currently, conservationists are concerned about the impact habitat fragmentation may have on the long-term persistence of the forest-dwelling guiña (Leopardus guigna), given the increasingly modified landscapes in which they live. Here we evaluate the effect habitat cover, fragmentation and anthropogenic pressure have on the occupancy probability for guiñas in privately-owned forest fragments. We collected camera-trap data from 100 temperate rainforest sites in Chile and used single-season occupancy modeling to evaluate the influence of 13 parameters of landscape structure/anthropogenic pressure and four parameters of detection probability on the ocurrence of guiñas. The camera-trap survey data comprised 4168 camera-trap days and 112 independent records of guiñas. Surprisingly, fragmented (defined as having a high perimeter-to-area ratio) and moderately sized habitat patches best predicted site occupancy. Occupancy also increased where habitat patches were closer to continuous forest and nearer to buildings. Our results imply that guiñas can benefit from a high degree of edge type habitats in fragmented landscapes, capable of adapting to habitat fragmentation in the proximity to large continuous forest patches. This suggests that guiñas have a broader niche than previously believed. Additionally, the guiña is tolerant of human infrastructure. Further research is required to identify potential ecological traps, long-term source-sink dynamics, and the habitat loss/fragmentation threshold beyond which guiña populations are no longer viable.     

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。     

Temperature can interact with landscape factors to affect songbird productivity

Increased temperatures and more extreme weather patterns associated with global climate change can interact with other factors that regulate animal populations, but many climate change studies do not incorporate other threats to wildlife in their analyses. We used 20 years of nest‐monitoring data from study sites across a gradient of habitat fragmentation in Missouri, USA, to investigate the relative influence of weather variables (temperature and precipitation) and landscape factors (forest cover and edge density) on the number of young produced per nest attempt (i.e., productivity) for three species of songbirds. We detected a strong forest cover × temperature interaction for the Acadian Flycatcher (Empidonax virescens) on productivity. Greater forest cover resulted in greater productivity because of reduced brood parasitism and increased nest survival, whereas greater temperatures reduced productivity in highly forested landscapes because of increased nest predation but had no effect in less forested landscapes. The Indigo Bunting (Passerina cyanea) exhibited a similar pattern, albeit with a marginal forest cover × temperature interaction. By contrast, productivity of the Northern Cardinal (Cardinalis cardinalis) was not influenced by landscape effects or temperature. Our results highlight a potential difficulty of managing wildlife in response to global change such as habitat fragmentation and climate warming, as the habitat associated with the greatest productivity for flycatchers was also that most negatively influenced by high temperatures. The influence of high temperatures on nest predation (and therefore, nest predators) underscores the need to acknowledge the potential complexity of species' responses to climate change by incorporating a more thorough consideration of community ecology in the development of models of climate impacts on wildlife.     

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.     

Under‐representation of avian studies in landscape genetics

Landscape genetics is a rapidly growing discipline that examines how heterogeneous landscapes and other environmental factors influence population genetic variation. We conducted a systematic review of the landscape genetic literature which demonstrates that birds are severely under‐represented relative to their species diversity and general publication prevalence. Most avian studies were on species that have relatively low dispersal ability, and we suggest that this reflects an assumed high vagility of birds that precludes spatial genetic variation relatable to landscape heterogeneity. However, spatial genetic variation exists in several bird species with very high dispersal ability, but this has not been considered in the context of landscape features. Genetic patterns may also relate to landscape due to breeding habitat selection and territorial behaviour, despite the fact that species may be able to move throughout different landscape elements with minimal movement costs. Habitat loss and fragmentation are continuing globally and are strongly related to declines in bird populations. Landscape genetic studies provide a means to understand, predict and mitigate the effects of anthropogenic landscape change on birds. This review promotes the need for landscape genetic studies of birds, such that a greater understanding of the drivers of their genetic structuring can be developed and generalizations can be made from landscape genetic studies that apply more broadly across taxa.     

Developing a functional connectivity indicator to detect change in fragmented landscapes

Biodiversity indicators are increasingly used to assess progress towards conservation targets. Particular indicators are required to assess the impacts of habitat fragmentation on landscape connectivity and biodiversity value. This paper recognises that connectivity is best defined by the interaction between species and the landscape in which they occur, and proposes a functional approach to assess connectivity. The approach utilises an incidence function model (IFM) as a spatially explicit method to assess potential species-level connectivity. The standard IFM connectivity measure is modified to account for the influence of the surrounding landscape matrix on edge impacts (through a weighted internal edge buffer) and ecological isolation (through an assessment of least-cost distance to account for landscape permeability). It has been recognised that such patch-based connectivity measures can provide misleading results when used to examine change, as they only focus on between patch movements. As a result, a modified hybrid IFM, based on a combination of patch and cell-based approaches, is developed to account for both within (intra) and between (inter) patch connectivity. The resulting probability of functional connectivity (PFC) indicator was evaluated, alongside a patch-based connectivity measure, through the application to four model landscapes based on changes (2 negative and 2 positive) to a control landscape. The four model landscapes illustrate the impact of landscape change on habitat area, edge impacts and matrix permeability. The proposed PFC indicator successfully discriminated between the two negative and the two positive changes to the control landscape, whereas, the patch-based connectivity measure detected change successfully within three of the four landscapes. The PFC indicator predicted a decrease in intra and inter-patch connectivity following habitat loss and fragmentation (negative change 1), whereas patch-based connectivity measures indicate an increase in connectivity between fragmented patches. The proposed PFC indicator offers the opportunity to take the necessary species-based perspective to examine functional connectivity, incorporating habitat preference, dispersal probability, edge impacts and ecological isolation/permeability. The urgency to assess changes in connectivity and support conservation policy means that there is little time to wait for more complete data. We believe the proposed approach provides a robust balance between the data required and the biologically meaningful indicator produced.     

Matrix quality and disturbance frequency drive evolution of species behavior at habitat boundaries

Previous theoretical studies suggest that a species' landscape should influence the evolution of its dispersal characteristics, because landscape structure affects the costs and benefits of dispersal. However, these studies have not considered the evolution of boundary crossing, that is, the tendency of animals to cross from habitat to nonhabitat (“matrix”). It is important to understand this dispersal behavior, because of its effects on the probability of population persistence. Boundary‐crossing behavior drives the rate of interaction with matrix, and thus, it influences the rate of movement among populations and the risk of dispersal mortality. We used an individual‐based, spatially explicit model to simulate the evolution of boundary crossing in response to landscape structure. Our simulations predict higher evolved probabilities of boundary crossing in landscapes with more habitat, less fragmented habitat, higher‐quality matrix, and more frequent disturbances (i.e., fewer generations between local population extinction events). Unexpectedly, our simulations also suggest that matrix quality and disturbance frequency have much stronger effects on the evolution of boundary crossing than either habitat amount or habitat fragmentation. Our results suggest that boundary‐crossing responses are most affected by the costs of dispersal through matrix and the benefits of escaping local extinction events. Evolution of optimal behavior at habitat boundaries in response to the landscape may have implications for species in human‐altered landscapes, because this behavior may become suboptimal if the landscape changes faster than the species' evolutionary response to that change. Understanding how matrix quality and habitat disturbance drive evolution of behavior at boundaries, and how this in turn influences the extinction risk of species in human‐altered landscapes should help us identify species of conservation concern and target them for management.     

Interactive effects of landscape and weather on dispersal

Over the last decades, many species have been forced to track their shifting climate envelopes, and at the same time man‐induced landscape fragmentation has led to the global decrease of natural habitat availability and connectivity. The interaction between these two co‐occurring global environmental changes might have very strong effects on biodiversity that are still understudied. Species‐specific responses to these environmental changes critically depend on individual dispersal, either to track suitable climatic conditions or to cope with landscape fragmentation. Here we study how dispersal in an ectotherm is affected by interactions between landscape fragmentation and weather conditions. We show that both the emigration rates out of suitable habitats and the topology of the trajectory of dispersing individuals were affected by temperature and landscape fragmentation. The emigration rate was temperature‐dependent in fragmented landscapes, with butterflies emigrating more at high temperatures. The emigration rate was temperature insensitive in more continuous landscapes. Move length was farther at low temperatures and less at high temperatures in fragmented landscapes. Move length was less at low temperatures and farther at high temperatures in more continuous landscapes. To our knowledge only two recent studies have documented patterns of interactions between climate and fragmentation, despite the fact that they are the two main drivers of biodiversity loss worldwide. Here, we go a step further by providing mechanistic explanations to such patterns.