Context

Anthropogenic activities readily result in the fragmentation of habitats such that species persistence increasingly depends on their ability to disperse. However, landscape features that enhance or limit individual dispersal are often poorly understood. Landscape genetics has recently provided innovative solutions to evaluate landscape resistance to dispersal.

Objectives

We studied the dispersal of the common meadow brown butterfly, Maniola jurtina, in agricultural landscapes, using a replicated study design and rigorous statistical analyses. Based on existing behavioral and life history research, we hypothesized that the meadow brown would preferentially disperse through its preferred grassy habitats (meadows and road verges) and avoid dispersing through woodlands and the agricultural matrix.

Methods

Samples were collected in 18 study landscapes of 5 × 5 km in three contrasting agricultural French regions. Using circuit theory, least cost path and transect-based methods, we analyzed the effect of the landscape on gene flow separately for each sex.

Results

Analysis of 1681 samples with 6 microsatellites loci revealed that landscape features weakly influence meadow brown butterfly gene flow. Gene flow in both sexes appeared to be weakly limited by forests and arable lands, whereas grasslands and grassy linear elements (road verges) were more likely to enhance gene flow.

Conclusion

Our results are consistent with the hypothesis of greater dispersal through landscape elements that are most similar to suitable habitat. Our spatially replicated landscape genetics study allowed us to detect subtle landscape effects on butterfly gene flow, and these findings were reinforced by consistent results across analytical methods.

Context

The relative importance of habitat area and connectivity for species richness is often unknown. Connectivity effects may be confounded with area effects or they may be of minor importance as posited by the habitat-amount hypothesis.

Objectives

We studied effects of habitat area and connectivity of linear landscape elements for plant species richness at plot level. We hypothesized that connectivity of linear landscape elements, assessed by resistance distance, has a positive effect on species richness beyond the effect of area and, further, that the relative importance of connectivity varies among groups of species with different habitat preferences and dispersal syndromes.

Methods

We surveyed plant species richness in 50 plots (25 m²) located on open linear landscape elements (field margins, ditches) in eight study areas of 1 km² in agricultural landscapes of Northwest Germany. We calculated the area of linear landscape elements and assessed their connectivity using resistance distance within circular buffers (500 m) around the plots. Effects of area and connectivity on species richness were modelled with generalised linear mixed models.

Results

Species richness did not increase with area. Resistance distance had significant negative effects on total richness and on the richness of typical species of grasslands and wetlands. Regarding dispersal syndromes, resistance distance had negative effects on the richness of species with short-distance, long-distance and aquatic dispersal. The significant effects of resistance distance indicated that species richness increased with connectivity of the network of linear landscape elements.

Conclusions

Connectivity is more important for plant species richness in linear landscape elements than area. In particular, the richness of plant species that are dispersal limited and confined to semi-natural habitats benefits from connective networks of linear landscape elements in agricultural landscapes.

Context

In deserts, many plant species exhibit a patchy spatial distribution within a harsh habitat matrix, where the likelihood of propagule dispersal among patches is uncertain, but may be promoted by landscape corridors or dispersal vectors.

Objectives

We examine the connectivity of a representative desert plant species (Acacia (Senegalia) greggii), and the ability of three major factors (animal dispersal agents, water flow along dry-washes, and climate) to facilitate dispersal within four watersheds in the Mojave National Preserve.

Methods

We genotyped 323 individuals sampled across 22 one-hectare sites using ten nuclear microsatellite markers.

Results

A hierarchical AMOVA revealed no significant differentiation among watersheds (F _RT = 0.00, P > 0.10), and very little genetic structure among all sites (F _ST = 0.03, P < 0.001), indicating regional connectivity. Mantel tests indicated distance along dry-washes best explained genetic distance between sites (r = 0.47, P < 0.05) when compared to Euclidean distance (P > 0.05), a distance measure based on rodent dispersal (P > 0.05), and a distance measure avoiding inhospitable climate (P > 0.05). An AIC comparison of generalized linear models found that within site genetic diversity (H _E and allelic richness) and average relatedness were best explained by slope (which increases seed dispersal potential via water flow) and area of the upstream watershed (which determines the number of potential seed donors), rather than plant density or habitat suitability.

Conclusions

Together, these findings indicate that dry-washes are key landscape features that enhance dispersal and regional connectivity in this patchy desert plant.

Context

Barriers to dispersal influence the ability of individuals to expand into new areas and can ultimately define success of reintroduction programs. American marten (Martes americana) were reintroduced to the Upper Peninsula of Michigan, USA, from multiple, genetically differentiated source populations from 1955 to 1992. Previous research found multiple genetic clusters near release sites with little admixing, suggesting barriers to dispersal exist.

Objectives

We sought to identify whether the contact zones between genetic clusters coincided with landscape features hypothesized to influence M. americana dispersal. We also investigated whether the degree of landscape contiguity within each genetic cluster differed among clusters.

Methods

We mapped cluster boundaries in M. americana genetic assignment probabilities and used correlation length as a measure of landscape contiguity between genetic clusters. We then evaluated the effects of land cover and roads on spatial genetic structure using a spatial autoregressive model.

Results

We found that gene flow was facilitated by contiguous coniferous forest and low incidence of roads. However, the strength of those relationships varied by genetic cluster. Contact zones among some genetic clusters spatially coincided with areas of high road and low conifer contiguity, compared to within-clusters.

Conclusions

In contrast to landscape genetic analyses focused on identifying barriers to gene flow, we incorporated methods that are relatively novel in landscape genetics to quantify how landscape contiguity influences spatial genetic structure. Using this method we were able to identify landscape barriers to dispersal at the genetic cluster boundaries for a reintroduced species distributed continuously across the landscape.

Context

Amphibian metapopulations have become increasingly fragmented in the Midwestern United States, with wetland-breeding salamanders being especially dependent on intact, high-quality forested landscapes. However, the degree to which amphibian populations are isolated, the factors that influence dispersal and, ultimately, functional connectivity remain areas in need of investigation.

Objectives and methods

We combined population demographic and genetic approaches to assess how a landscape fragmented by agriculture influences functional connectivity and metapopulation dynamics of a locally threatened salamander (Ambystoma jeffersonianum).

Results

We found that the allelic richness and heterozygosity of this species was significantly related to the level of connectivity with other occupied breeding wetlands and that decreased connectivity resulted in increased genetic differentiation. We also found that effective population size appears to be declining and, while correlative, our focal landscape has experienced significant losses of forested upland habitats and potential wetland breeding habitats over the last 200 years.

Conclusions

By combining population and landscape genetic analyses with an assessment of regional wetland occupancy, our study has uniquely synthesized genetic and metapopulation processes, while also incorporating the effects of the landscape matrix on dispersal, connectivity, and population differentiation. The significant relationship between connectivity with heterozygosity, allelic richness, and genetic divergence observed in this study reinforces empirical observations of long distance dispersal and movements in ambystomatid salamanders. However, our results show that protection of core habitat around isolated wetlands may not sufficiently minimize genetic differentiation among populations and preserve critical genetic diversity that may be essential for the long-term persistence of local populations.

Context

Methods for measuring restoration success that include functional connectivity between species’ populations are rare in landscape ecology and restoration practices. We developed an approach that analyzes connectivity between populations of target species and their dispersal probabilities to assess restoration success based on easily accessible input data. Applying this method to landscape development scenarios can help optimize restoration planning.

Objectives

We developed an assessment for restoration success and restoration planning based on functional connectivity between species’ populations and spatially explicit scenarios. The method was used in a case study to test its applicability.

Methods

Based on data on available habitat, species’ occurrence and dispersal ranges, connectivity metrics and dispersal probabilities for target species are calculated using the software Conefor Sensinode. The metrics are calculated for scenarios that reflect possible changes in the landscape to provide a basis for future restoration planning. We applied this approach to floodplain meadows along the Upper Rhine for four plant species and three future scenarios.

Results

In the case study, habitats of the target species were poorly connected. Peucedanum officinale and Sanguisorba officinalis were more successful in recolonizing new habitats than Iris spuria and Serratula tinctoria. The scenarios showed that restoration of species-rich grassland was beneficial for dispersal of the target species. As expected in the agriculturally dominated study area, restoration of former arable land significantly increased dispersal probabilities.

Conclusions

In the case study, the developed approach was easily applicable and provided reasonable results. Its implementation will be helpful in decision-making for future restoration planning.

Context

Quantifying gene flow in natural populations is a key topic in both evolutionary and conservation biology. Understanding the extent to which the landscape matrix facilitates or impedes gene flow is becoming a high priority in a context of worldwide habitat loss and fragmentation.

Objectives

Unexpectedly, a lower genetic diversity and a higher genetic structure have been previously observed in the less fragmented and the most forested habitat across four pine marten (Martes martes) populations in France. Our aim was to quantify the effect of landscape on the spatial distribution of genetic diversity in two populations in contrasting habitats.

Methods

We conducted an individual-based landscape genetics analysis in a highly fragmented rural plain (Bresse, n = 126) and in a highly forested (50 %) mountainous area (Ariège, n = 88) in France. We tested for isolation-by-resistance using least-cost distances and used a causal modeling approach on 16,384 landscape and 104 elevation resistance scenarios.

Results

Landscape structure influenced the genetic differentiation in Bresse, with vegetation providing more genetic connectivity over the study area than open areas, while roads and human buildings showed unexpected low resistance to gene flow. In Ariège, genetic differentiation was mainly associated with changes in elevation, with an optimal elevation for gene flow of around 1700 m, likely associated with changes in vegetation structure.

Conclusions

The pine marten seems to be able to cope with human-dominated landscapes and with fragmented forest landscapes, whereas elevation is the major driver of genetic differentiation in our mountainous landscape. Additionally, we highlight the importance of spatial replication in landscape genetics for deriving reliable conservation and management measures over the species distribution.

Context

Various species of forest trees are commonly used for ornamental purposes and are therefore frequently found in nonforest ecosystems. They constitute an important component of the so-called trees outside forests (TOF). Not much is known, however, about the drivers of TOF spatial distribution either in urbanized or in agricultural landscapes since they are generally absent from forest inventories.

Objective

The present study focused on the spatial distribution of TOF across agricultural landscapes and their potential role in the dispersal of a forest pest insect, the pine processionary moth, Thaumetopoea pityocampa (PPM).

Methods

All the TOF belonging to the genera Pinus, Cedrus and Pseudotsuga were considered as potential hosts and inventoried within a 22 × 22 km study window. We fitted a nonstationary Poisson process to the empirical data and used the distance to the nearest building as a covariate.

Results

Both empirical and simulated data indicated that TOF associated to human artifacts/urbanized areas constituted the main source of landscape connectivity for the PPM in the open fields under study. Because they do not account for TOF, forest inventories dramatically underestimate landscape connectivity and provide an erroneous picture of the PPM habitat distribution.

Conclusions

We conclude that TOF, especially the ornamental component, must be taken into account when it comes to understanding forest insect landscape dynamics or genetics. The omnipresence of TOF also suggests a potentially huge role in pest dispersal and invasive species expansion.

Context

Seed dispersal is recognized as having profound effects on the distribution, dynamics and structure of plant populations and communities. However, knowledge of how landscape structure shapes carnivore-mediated seed dispersal patterns is still scarce, thereby limiting our understanding of large-scale plant population processes.

Objectives

We aim to determine how the amount and spatial configuration of forest cover impacted the relative abundance of carnivorous mammals, and how these effects cascaded through the seed dispersal kernels they generated.

Methods

Camera traps activated by animal movement were used for carnivore sampling. Colour-coded seed mimics embedded in common figs were used to know the exact origin of the dispersed seed mimics later found in carnivore scats. We applied this procedure in two sites differing in landscape structure.

Results

We did not find between-site differences in the relative abundance of the principal carnivore species contributing to seed dispersal patterns, Martes foina. Mean dispersal distance and the probability of long dispersal events were higher in the site with spatially continuous and abundant forest cover, compared to the site with spatially aggregated and scarcer forest cover. Seed deposition closely matched the spatial patterning of forest cover in both study sites, suggesting behaviour-based mechanisms underpinning seed dispersal patterns generated by individual frugivore species.

Conclusions

Our results provide the first empirical evidence of the impact of landscape structure on carnivore-mediated seed dispersal kernels. They also indicate that seed dispersal kernels generated strongly depend on the effect that landscape structure exerts on carnivore populations, particularly on habitat-use preferences.

Context

Many aquatic communities are linked by the aerial dispersal of multiple, interacting species and are thus structured by processes occurring in both the aquatic and terrestrial compartments of the ecosystem.

Objectives

To evaluate the environmental factors shaping the aquatic macroinvertebrate communities associated with tank bromeliads in an urban landscape.

Methods

Thirty-two bromeliads were georeferenced to assess the spatial distribution of the aquatic meta-habitat in one city. The relative influence of the aquatic and terrestrial habitats on the structure of macroinvertebrate communities was analyzed at four spatial scales (radius = 10, 30, 50, and 70 m) using redundancy analyses.

Results

We sorted 18,352 aquatic macroinvertebrates into 29 taxa. Water volume and the amount of organic matter explained a significant part of the taxa variance, regardless of spatial scale. The remaining variance was explained by the meta-habitat size (i.e., the water volume for all of the bromeliads within a given surface area), the distance to the nearest building at small scales, and the surface area of buildings plus ground cover at larger scales. At small scales, the meta-habitat size influenced the two most frequent mosquito species in opposite ways, suggesting spatial competition and coexistence. Greater vegetation cover favored the presence of a top predator.

Conclusions

The size of the meta-habitat and urban landscape characteristics influence the structure of aquatic communities in tank bromeliads, including mosquito larval abundance. Modifications to this landscape will affect both the terrestrial and aquatic compartments of the urban ecosystem, offering prospects for mosquito management during urban planning.

Context

Scale dependence of bat habitat selection is poorly known with few studies evaluating relationships among landscape metrics such as class versus landscape, or metrics that measure composition or configuration. This knowledge can inform conservation approaches to mitigate habitat loss and fragmentation.

Objectives

We evaluated scale dependence of habitat associations and scaling patterns of landscape metrics in relation to bat occurrence or capture rate in forests of southwestern Nicaragua.

Methods

We captured 1537 bats at 35 locations and measured landscape and class metrics across 10 spatial scales (100–1000 m) surrounding capture locations. We conducted univariate scaling across the 10 scales and identified scales and variables most related to bat occurrence or capture rate.

Results

Edge and patch density, at both landscape and class levels, were the most important variables across species. Feeding guilds varied in their response to metrics. Certain landscape and configuration metrics were most influential at fine (100 m) and/or broad (1000 m) spatial scales while most class and composition metrics were influential at intermediate scales.

Conclusions

These results provide insight into the scale dependence of habitat associations of bat species and the influence of fine and broad scales on habitat associations. The effects of scale, examined in our study and others from fine (100 m) to broad (5 km) indicate habitat relationships for bats may be more informative at larger scales. Our results suggest there could be general differences in scale relationships for different groups of landscape metrics, which deserves further evaluation in other taxonomic groups.

Context

Landscape heterogeneity (the composition and configuration of different landcover types) plays a key role in shaping woodland bird assemblages in wooded-agricultural mosaics. Understanding how species respond to landscape factors could contribute to preventing further decline of woodland bird populations.

Objective

To investigate how woodland birds with different species traits respond to landscape heterogeneity, and to identify whether specific landcover types are important for maintaining diverse populations in wooded-agricultural environments.

Methods

Birds were sampled from woodlands in 58 2 × 2 km tetrads across southern Britain. Landscape heterogeneity was quantified for each tetrad. Bird assemblage response was determined using redundancy analysis combined with variation partitioning and response trait analyses.

Results

For woodland bird assemblages, the independent explanatory importance of landscape composition and landscape configuration variables were closely interrelated. When considered simultaneously during variation partitioning, the community response was better represented by compositional variables. Different species responded to different landscape features and this could be explained by traits relating to woodland association, foraging strata and nest location. Ubiquitous, generalist species, many of which were hole-nesters or ground foragers, correlated positively with urban landcover while specialists of broadleaved woodland avoided landscapes containing urban areas. Species typical of coniferous woodland correlated with large conifer plantations.

Conclusions

At the 2 × 2 km scale, there was evidence that the availability of resources provided by proximate landcover types was highly important for shaping woodland bird assemblages. Further research to disentangle the effects of composition and configuration at different spatial scales is advocated.

Context

With accelerated land-use change throughout the world, increased understanding of the relative effects of landscape composition and configuration on biological system and bioinvasion in particular, is needed to design effective management strategies. However, this topic is poorly understood in part because empirical studies often fail to account for large gradients of habitat complexity and offer insufficient or even no replication across habitats.

Objectives

The aim of this study was to disentangle the independent and interactive effects of landscape composition and landscape configuration on the establishment and spread of invasive insect species.

Methods

We explore a spatially-explicit, mechanistic modeling framework that allows for systematic investigation of the impact of changes in landscape composition and landscape configuration on establishment and spread of invasive insect species. Landscape metrics are used as an indicators of invasive insect establishment and spread.

Results

We showed that the presence of an Allee effect leads to a balance between the effectiveness of spread and invasion success. Spread is maximized at an intermediate dispersal level and inhibited at both low and high levels of dispersal. The landscape, by either increasing or mitigating the dispersal abilities of a species, can lead to a rate of spread under a dispersal threshold for which density and spread is at the highest.

Conclusion

Our study proposes that change in landscape structure is an additional explanation of the highly variable spread dynamics observed in natural and anthropogenic landscapes. Consequently, a landscape-scale perspective could significantly improve spread risk assessment and the design of control or containment strategies.

Context

Habitat fragmentation generates a loss of functional connectivity detrimental to the persistence of biodiversity. The French agricultural intensification initiated in the 1950s has caused a decline in field margins.

Objectives

As field margins may facilitate species dispersal while providing socio-economic benefits, it is of interest to assess their contribution to the functional connectivity of insect-pollinated plants in agro-ecosystems. This will help develop appropriate management strategies mitigating fragmentation.

Methods

We addressed this issue by studying the links between landscape structure and the patterns of abundance and pollen dispersal (using fluorescent dye particles) for two contrasted insect-pollinated plants occurring in field margins (Crepis sancta and Euphorbia serrata). We investigated the influence of field margins quality and of the surrounding matrix on pollen dispersal and compared the relevance of the least-cost algorithm with a straight-line approach to depict pollinators’ movements.

Results

The influence of landscape structure on plant abundance is species and scale-specific. Pollen dispersal decreases with distance from the source. For E. serrata, it was preferentially dispersed via field margins, confirming the relevance of the least-cost algorithm, while C. sancta dispersal followed a straight-line.

Conclusions

Euphorbia serrata, which grows strictly on field margins with a greater dispersal ability and a more diversified pollinator guild than C. sancta, is less affected by land-use changes. Our study demonstrates the contrasting contributions of field margins to pollen dispersal as they may act as functional corridors favouring pollinators’ movement depending on the species of interest.

Context

Landscape fragmentation significantly affects species distributions by decreasing the number and connectivity of suitable patches. While researchers have hypothesized that species functional traits could help in predicting species distribution in a landscape, predictions should depend on the type of patches available and on the ability of species to disperse and grow there.

Objectives

To explore whether different traits can explain the frequency of grassland species (number of occupied patches) and/or their occupancy (ratio of occupied to suitable patches) across a variety of patch types within a fragmented landscape.

Methods

We sampled species distributions over 1300 grassland patches in a fragmented landscape of 385 km² in the Czech Republic. Relationships between functional traits and species frequency and occupancy were tested across all patches in the landscape, as well as within patches that shared similar management, wetness, and isolation.

Results

Although some traits predicting species frequency also predicted occupancy, others were markedly different, with competition- and dispersal-related traits becoming more important for occupancy. Which traits were important differed for frequency and occupancy and also differed depending on patch management, wetness, and isolation.

Conclusions

Plant traits can provide insight into plant distribution in fragmented landscapes and can reveal specific abiotic, biotic, and dispersal processes affecting species occurrence in a patch type. However, the importance of individual traits depends on the type of suitable patches available within the landscape.

Context

The problem of how ecological mechanisms create and interact with patterns across different scales is fundamental not only for understanding ecological processes, but also for interpretations of ecological dynamics and the strategies that organisms adopt to cope with variability and cross-scale influences.

Objectives

Our objective was to determine the consistency of the role of individual habitat patches in pattern-process relationships (focusing on the potential for dispersal within a network of patches in a fragmented landscape) across a range of scales.

Methods

Network analysis was used to assess and compare the potential connectivity and spatial distribution of highland fynbos habitat in and between protected areas of the Western Cape of South Africa. Connectivity of fynbos patches was measured using ten maximum threshold distances, ranging from five to 50 km, based on the known average dispersal distances of fynbos endemic bird species.

Results

Network connectivity increased predictably with scale. More interestingly, however, the relative contributions of individual protected areas to network connectivity showed strong scale dependence.

Conclusions

Conservation approaches that rely on single-scale analyses of connectivity and context (e.g., based on data for a single species with a given dispersal distance) are inadequate to identify key land parcels. Landscape planning, and specifically the assessment of the value of individual areas for dispersal, must therefore be undertaken with a multi-scale approach. Developing a better understanding of scaling dependencies in fragmenting landscapes is of high importance for both ecological theory and conservation planning.

Context

A challenge devising revegetation strategies in fragmented landscapes is conserving for the widest spectrum of biodiversity. Habitat network reconstruction should improve landscape capacity to maintain species populations. However, the location of revegetation often fails to account for species occurrence and dispersal processes operating across spatial scales.

Objectives

Our objective was to integrate metapopulation theory with estimates of landscape capacity and dispersal pathways to highlight connectivity gaps. Maintenance of populations could thereby be facilitated through reconnecting habitat networks across regional and broader scales, with assumed benefit for the dispersal needs of less sensitive species.

Methods

Predicted occupancy and metapopulation capacity were calculated for a generic focal species derived from fragmentation-sensitive woodland birds, mammals and reptiles. A metapopulation connectivity analysis predicted regional dispersal links to identify likely routes through which individuals may move to contribute to the viability of the population. We used the revegetation programmes of the Brigalow–Nandewar Biolinks project, eastern New South Wales, Australia, to demonstrate our approach.

Results

Landscape capacity of the current landscape varied across the region. Low-value links between populations provided greatest opportunities for revegetation and improved landscape capacity. Where regional connectivity did not indicate a pathway between populations, broader scale connectivity provided guidance for revegetation.

Conclusions

The metapopulation-based model, coupled with a habitat dispersal network analysis, provided a platform to inform revegetation locations and better support biodiversity. Our approach has application for directing on-ground action to support viable populations, assess the impact of revegetation schemes or monitor the progress of staged implementations.

Context

Corridors are usually delineated as areas of minimum cumulative resistance to movement through a resistance surface and characterized by their effective distance (accumulated resistance along the least-cost path). The results of these assessments depend on resistance values, which are typically derived from the inverse of habitat suitability models or from presence data of individuals within their home ranges, rather than from data on dispersal or exploratory movements.

Objective

Evaluate the extent to which corridor delineation and effective distance estimates may vary depending on whether home range locations or dispersal data are used to characterize species habitat selection and landscape resistance to movement.

Methods

We analyzed a large telemetry dataset (GPS collars) for the endangered Iberian lynx. We modeled corridors and effective distances three ways: (1) considering only GPS locations within home ranges, (2) considering only locations in dispersal or exploratory movements outside home ranges, and (3) considering all locations together.

Results

Delineated least-cost corridors followed similar trajectories and sometimes overlapped in the three models. The estimated effective distances were 42 % lower in the dispersal-based model than in the model based solely on home range use.

Conclusions

Models derived exclusively from locations within home ranges may provide lower connectivity estimates than models derived from dispersal locations, affecting estimates of resistance to move between habitat areas, even when the most likely movement routes are similar. Although dispersal data are costly to gather, they potentially provide more realistic assessments of the actual isolation of populations in heterogeneous landscapes.

Context

The abundance of important providers of ecosystem services such as wild bees likely increases with landscape heterogeneity, but may also fluctuate across the flowering season following varying weather conditions.

Objectives

In the present study, we investigated the combined effect of landscape heterogeneity and intra-annual variability in temperature and precipitation on the spatial and temporal stability of wild bee abundance.

Methods

We used bee monitoring data from six 4 km × 4 km sites in central Germany and 16 local communities per site. The data were collected six times per year from 2010 to 2013. Following a multimodel inference approach, we identified the importance of landscape heterogeneity, weather variability and their interaction to the stability of wild bee abundance.

Results

We found that the stability of wild bee abundance increased with landscape heterogeneity, but decreased with increasing intra-annual variability in both temperature and precipitation. However, our key finding was a buffering mechanism enabling high abundance stability in heterogeneous landscapes even under highly variable temperature conditions. Interestingly, the same mechanism did not apply for high variability in precipitation.

Conclusions

Our findings suggest that increasing landscape heterogeneity is beneficial for protecting wild bees against the projected increase in temperature variability until the end of the twenty first century, although we cannot make inferences for extreme events such as heatwaves. Nevertheless, our results equally highlight that landscape heterogeneity should not be treated as a one-size-fits-all solution and the need remains for developing alternative strategies to mitigate the effect of increasing variability in precipitation.