Where and why are species' range shifts hampered by unsuitable landscapes?

There is widespread concern that species will fail to track climate change if habitat is too scarce or insufficiently connected. Targeted restoration has been advocated to help species adapt, and a "conductance" metric has been proposed, based on simulation studies, to predict effective habitat configurations. However, until now there is very little empirical evidence on how the configuration of habitat is affecting expansion at species' cool range margins. We analysed the colonisation events that have occurred in continuously monitored trap locations for 54 species of southerly distributed moths in Britain between 1985 and 2011. We tested whether the time until colonisation was affected by attributes of each species, and of intervening landcover and climate between the trap and the baseline distribution (1965-1985). For woodland species, the time until colonisation of new locations was predicted by the "conductance" of woodland habitat, and this relationship was general, regardless of species' exact dispersal distances and habitat needs. This shows that contemporary range shifts are being influenced by habitat configuration as well as simple habitat extent. For species associated with farmland or suburban habitats, colonisation was significantly slower through landscapes with a high variance in elevation and/or temperature. Therefore, it is not safe to assume that such relatively tolerant species face no geographical barriers to range expansion. We thus elucidate how species' attributes interact with landscape characteristics to create highly heterogeneous patterns of shifting at cool range margins. Conductance, and other predictors of range shifts, can provide a foundation for developing coherent conservation strategies to manage range shifts for entire communities.

Biodiversity and yield trade-offs for organic farming.

Organic farming supports higher biodiversity than conventional farming, but at the cost of lower yields. We conducted a meta-analysis quantifying the trade-off between biodiversity and yield, comparing conventional and organic farming. We developed a compatibility index to assess whether biodiversity gains from organic farming exceed yield losses, and a substitution index to assess whether organic farming would increase biodiversity in an area if maintaining total production under organic farming would require cultivating more land at the expense of nature. Overall, organic farming had 23% gain in biodiversity with a similar cost of yield decline. Biodiversity gain is negatively correlated to yield loss for microbes and plants, but no correlation was found for other taxa. The biodiversity and yield trade-off varies under different contexts of organic farming. The overall compatibility index value was close to zero, with negative values for cereal crops, positive for non-cereal crops, and varies across taxa. Our results indicate that, on average, the proportion of biodiversity gain is similar to the proportion of yield loss for paired field studies. For some taxa in non-cereal crops, switching to organic farming can lead to a biodiversity gain without yield loss. We calculated the overall value of substitution index and further discussed the application of this index to evaluate when the biodiversity of less intensified farming system is advantageous.

Comparison between window traps and pan traps in monitoring flower-visiting insects in agricultural fields.

Sampling flower-visiting insects in agricultural fields at large spatial and temporal scales is significant for understanding local insect pollinator communities. The most commonly used method, pan trap, has been criticized due to its attractant bias. A window trap (also referred to as the flight-intercept trap) is a non-attractant sampling method, which has been applied in forests and grasslands, but rarely in agricultural fields. We aim to test whether we can replace pan traps with window traps in agricultural fields by comparing species richness and species composition between the two methods, and to show whether flower-visiting insects collected in both traps can reflect flower-visiting activity recorded by camera observation. We conducted a 2-year study to compare the performance of these sampling methods in an oilseed rape field. Results showed that the relative abundance of dominant flower-visiting species was highly correlated between the window trap and the pan trap samples, while window traps caught more individuals and higher (rarefied) species richness than pan traps. The species composition of window traps was more similar to each other than that of pan traps. The proportion of honey bees (Apis spp.) collected in both traps underestimated their flower-visiting activity recorded by camera observations, while sweat bees (Halictidae) and butterflies (Lepidoptera) were overestimated. Our study suggests that the window trap has the potential to serve as an alternative sampling method of flower-visiting insects to the pan trap. However, we need to be cautious when using specimens caught in both traps as a proxy of their flower-visiting activity.

Habitat patches providing south-north connectivity are under-protected in a fragmented landscape.

As species' ranges shift to track climate change, conservationists increasingly recognize the need to consider connectivity when designating protected areas (PAs). In fragmented landscapes, some habitat patches are more important than others in maintaining connectivity, and methods are needed for their identification. Here, using the Condatis methodology, we model range expansion through an adaptation of circuit theory. Specifically, we map 'flow' through 16 conservation priority habitat networks in England, quantifying how patches contribute to functional South-North connectivity. We also explore how much additional connectivity could be protected via a connectivity-led protection procedure. We find high-flow patches are often left out of existing PAs; across 12 of 16 habitat networks, connectivity protection falls short of area protection by 13.6% on average. We conclude that the legacy of past protection decisions has left habitat-specialist species vulnerable to climate change. This situation may be mirrored in many countries which have similar habitat protection principles. Addressing this requires specific planning tools that can account for the directions species may shift. Our connectivity-led reserve selection procedure efficiently identifies additional PAs that prioritize connectivity, protecting a median of 40.9% more connectivity in these landscapes with just a 10% increase in area.

Wing morphological responses to latitude and colonisation in a range expanding butterfly.

Populations undergoing rapid climate-driven range expansion experience distinct selection regimes dominated both by increased dispersal at the leading edges and steep environmental gradients. Characterisation of traits associated with such expansions provides insight into the selection pressures and evolutionary constraints that shape demographic and evolutionary responses. Here we investigate patterns in three components of wing morphology (size, shape, colour) often linked to dispersal ability and thermoregulation, along latitudinal gradients of range expansion in the Speckled Wood butterfly (Pararge aegeria) in Britain (two regions of expansion in England and Scotland). We measured 774 males from 54 sites spanning 799 km with a 10-year mean average temperature gradient of 4 °C. A geometric morphometric method was used to investigate variation in size and shape of forewings and hindwings; colour, pattern, and contrast of the wings were examined using a measure of lightness (inverse degree of melanism). Overall, wing size increased with latitude by ∼2% per 100 km, consistent with Bergmann's rule. Forewings became more rounded and hindwings more elongated with history of colonisation, possibly reflecting selection for increased dispersal ability. Contrary to thermal melanism expectations, wing colour was lighter where larvae developed at cooler temperatures and unrelated to long-term temperature. Changes in wing spot pattern were also detected. High heterogeneity in variance among sites for all of the traits studied may reflect evolutionary time-lags and genetic drift due to colonisation of new habitats. Our study suggests that temperature-sensitive plastic responses for size and colour interact with selection for dispersal traits (wing size and shape). Whilst the plastic and evolutionary responses may in some cases act antagonistically, the rapid expansion of P. aegeria implies an overall reinforcing effect between these two mechanisms.

Incorporating connectivity into conservation planning for the optimal representation of multiple species and ecosystem services.

Conservation planning tends to focus on protecting species' ranges or landscape connectivity but seldom both-particularly in the case of diverse taxonomic assemblages and multiple planning goals. Therefore, information on potential trade-offs between maintaining landscape connectivity and achieving other conservation objectives is lacking. We developed an optimization approach to prioritize the maximal protection of species' ranges, ecosystem types, and forest carbon stocks, while also including habitat connectivity for range-shifting species and dispersal corridors to link protected area. We applied our approach to Sabah, Malaysia, where the state government mandated an increase in protected-area coverage of approximately 305,000 ha but did not specify where new protected areas should be. Compared with a conservation planning approach that did not incorporate the 2 connectivity features, our approach increased the protection of dispersal corridors and elevational connectivity by 13% and 21%, respectively. Coverage of vertebrate and plant species' ranges and forest types were the same whether connectivity was included or excluded. Our approach protected 2% less forest carbon and 3% less butterfly range than when connectivity features were not included. Hence, the inclusion of connectivity into conservation planning can generate large increases in the protection of landscape connectivity with minimal loss of representation of other conservation targets.

Incorporación de la Conectividad a la Planeación de la Conservación para la Representación Óptima de Especies Múltiples y Servicios Ambientales Resumen Las tendencias de planeación de la conservación tienden a enfocarse en la protección de la distribución geográfica de las especies o en la conectividad de paisajes, pero rara vez se enfocan en ambas - particularmente para el caso de los ensamblajes taxonómicos y las metas múltiples de planeación. Por lo tanto, hay carencias en la información sobre las compensaciones potenciales entre mantener la conectividad de los paisajes y alcanzar otros objetivos de conservación. Desarrollamos una estrategia de optimización para priorizar la protección máxima de la distribución de las especies, los tipos de ecosistemas y los stocks de carbono de los bosques, a la vez que incluimos la conectividad del hábitat para las especies que modifican su distribución y los corredores de dispersión para conectar el área protegida. Aplicamos nuestra estrategia en Sabah, Malasia, en donde el gobierno estatal ordenó un incremento de ∼305, 000 ha en la cobertura de áreas protegidas sin especificar la ubicación de las nuevas áreas protegidas. En comparación con una estrategia de planeación de la conservación que no incorporó las dos características de la conectividad, nuestra estrategia incrementó la protección de los corredores de dispersión y la conectividad altitudinal en un 13% y 21% respectivamente. La cobertura de la distribución de las especies de plantas y vertebrados y de los tipos de bosque fue la misma con o sin la inclusión de la conectividad. Nuestra estrategia protegió 2% menos del carbono forestal y 3% menos de la distribución de mariposas que cuando no se incluyeron las características de conectividad en la estrategia. Por lo tanto, incluir a la conectividad en la planeación de la conservación puede generar grandes incrementos en la protección de la conectividad del paisaje con una pérdida mínima de representación para los demás objetivos de conservación.

Testing the benefits of conservation set-asides for improved habitat connectivity in tropical agricultural landscapes.

Habitat connectivity is important for tropical biodiversity conservation. Expansion of commodity crops, such as oil palm, fragments natural habitat areas, and strategies are needed to improve habitat connectivity in agricultural landscapes. The Roundtable on Sustainable Palm Oil (RSPO) voluntary certification system requires that growers identify and conserve forest patches identified as High Conservation Value Areas (HCVAs) before oil palm plantations can be certified as sustainable. We assessed the potential benefits of these conservation set-asides for forest connectivity.We mapped HCVAs and quantified their forest cover in 2015. To assess their contribution to forest connectivity, we modelled range expansion of forest-dependent populations with five dispersal abilities spanning those representative of poor dispersers (e.g. flightless insects) to more mobile species (e.g. large birds or bats) across 70 plantation landscapes in Borneo.Because only 21% of HCVA area was forested in 2015, these conservation set-asides currently provide few connectivity benefits. Compared to a scenario where HCVAs contain no forest (i.e. a no-RSPO scenario), current HCVAs improved connectivity by ~3% across all dispersal abilities. However, if HCVAs were fully reforested, then overall landscape connectivity could improve by ~16%. Reforestation of HCVAs had the greatest benefit for poor to intermediate dispersers (0.5-3 km per generation), generating landscapes that were up to 2.7 times better connected than landscapes without HCVAs. By contrast, connectivity benefits of HCVAs were low for highly mobile populations under current and reforestation scenarios, because range expansion of these populations was generally successful regardless of the amount of forest cover. Synthesis and applications. The Roundtable on Sustainable Palm Oil (RSPO) requires that High Conservation Value Areas (HCVAs) be set aside to conserve biodiversity, but HCVAs currently provide few connectivity benefits because they contain relatively little forest. However, reforested HCVAs have the potential to improve landscape connectivity for some forest species (e.g. winged insects), and we recommend active management by plantation companies to improve forest quality of degraded HCVAs (e.g. by enrichment planting). Future revisions to the RSPO's Principles and Criteria should also ensure that large (i.e. with a core area >2 km2) HCVAs are reconnected to continuous tracts of forest to maximize their connectivity benefits.

Faster movement in nonhabitat matrix promotes range shifts in heterogeneous landscapes.

Ecologists often assume that range expansion will be fastest in landscapes composed entirely of the highest-quality habitat. Theoretical models, however, show that range expansion depends on both habitat quality and habitat-specific movement rates. Using data from 78 species in 70 studies, we find that animals typically have faster movement through lower-quality environments (73% of published cases). Therefore, if we want to manage landscapes for range expansion, there is a trade-off between promoting movement with nonhostile matrix, and promoting population growth with high-quality habitat. We illustrate how this trade-off plays out with the use of an exemplar species, the Baltimore checkerspot butterfly. For this species, we calculate that the expected rate of range expansion is fastest in landscapes with ~15% high-quality habitat. Behavioral responses to nonhabitat matrix have often been documented in animal populations, but rarely included in empirical predictions of range expansion. Considering movement behavior could change land-planning priorities from focus on high-quality habitat only to integrating high- and low-quality land cover types, and evaluating the costs and benefits of different matrix land covers for range expansion.

Landscape simplification weakens the association between terrestrial producer and consumer diversity in Europe.

Land-use change is one of the primary drivers of species loss, yet little is known about its effect on other components of biodiversity that may be at risk. Here, we ask whether, and to what extent, landscape simplification, measured as the percentage of arable land in the landscape, disrupts the functional and phylogenetic association between primary producers and consumers. Across seven European regions, we inferred the potential associations (functional and phylogenetic) between host plants and butterflies in 561 seminatural grasslands. Local plant diversity showed a strong bottom-up effect on butterfly diversity in the most complex landscapes, but this effect disappeared in simple landscapes. The functional associations between plant and butterflies are, therefore, the results of processes that act not only locally but are also dependent on the surrounding landscape context. Similarly, landscape simplification reduced the phylogenetic congruence among host plants and butterflies indicating that closely related butterflies become more generalist in the resources used. These processes occurred without any detectable change in species richness of plants or butterflies along the gradient of arable land. The structural properties of ecosystems are experiencing substantial erosion, with potentially pervasive effects on ecosystem functions and future evolutionary trajectories. Loss of interacting species might trigger cascading extinction events and reduce the stability of trophic interactions, as well as influence the longer term resilience of ecosystem functions. This underscores a growing realization that species richness is a crude and insensitive metric and that both functional and phylogenetic associations, measured across multiple trophic levels, are likely to provide additional and deeper insights into the resilience of ecosystems and the functions they provide.

A trait-based approach for predicting species responses to environmental change from sparse data: how well might terrestrial mammals track climate change?

Estimating population spread rates across multiple species is vital for projecting biodiversity responses to climate change. A major challenge is to parameterise spread models for many species. We introduce an approach that addresses this challenge, coupling a trait-based analysis with spatial population modelling to project spread rates for 15 000 virtual mammals with life histories that reflect those seen in the real world. Covariances among life-history traits are estimated from an extensive terrestrial mammal data set using Bayesian inference. We elucidate the relative roles of different life-history traits in driving modelled spread rates, demonstrating that any one alone will be a poor predictor. We also estimate that around 30% of mammal species have potential spread rates slower than the global mean velocity of climate change. This novel trait-space-demographic modelling approach has broad applicability for tackling many key ecological questions for which we have the models but are hindered by data availability.

Refugia and connectivity sustain amphibian metapopulations afflicted by disease.

Metapopulation persistence in fragmented landscapes depends on habitat patches that can support resilient local populations and sufficient connectivity between patches. Yet epidemiological theory for metapopulations has largely overlooked the capacity of particular patches to act as refuges from disease, and has suggested that connectivity can undermine persistence. Here, we show that relatively warm and saline wetlands are environmental refuges from chytridiomycosis for an endangered Australian frog, and act jointly with connectivity to sustain frog metapopulations. We coupled models of microclimate and infection probability to map chytrid prevalence, and demonstrate a strong negative relationship between chytrid prevalence and the persistence of frog populations. Simulations confirm that frog metapopulations are likely to go extinct when they lack environmental refuges from disease and lose connectivity between patches. This study demonstrates that environmental heterogeneity can mediate host-pathogen interactions in fragmented landscapes, and provides evidence that connectivity principally supports host metapopulations afflicted by facultative pathogens.

Range expansion through fragmented landscapes under a variable climate.

Ecological responses to climate change may depend on complex patterns of variability in weather and local microclimate that overlay global increases in mean temperature. Here, we show that high-resolution temporal and spatial variability in temperature drives the dynamics of range expansion for an exemplar species, the butterfly Hesperia comma. Using fine-resolution (5 m) models of vegetation surface microclimate, we estimate the thermal suitability of 906 habitat patches at the species' range margin for 27 years. Population and metapopulation models that incorporate this dynamic microclimate surface improve predictions of observed annual changes to population density and patch occupancy dynamics during the species' range expansion from 1982 to 2009. Our findings reveal how fine-scale, short-term environmental variability drives rates and patterns of range expansion through spatially localised, intermittent episodes of expansion and contraction. Incorporating dynamic microclimates can thus improve models of species range shifts at spatial and temporal scales relevant to conservation interventions.

The speed of range shifts in fragmented landscapes.

Species may be driven extinct by climate change, unless their populations are able to shift fast enough to track regions of suitable climate. Shifting will be faster as the proportion of suitable habitat in the landscape increases. However, it is not known how the spatial arrangement of habitat will affect the speed of range advance, especially when habitat is scarce, as is the case for many specialist species. We develop methods for calculating the speed of advance that are appropriate for highly fragmented, stochastic systems. We reveal that spatial aggregation of habitat tends to reduce the speed of advance throughout a wide range of species parameters: different dispersal distances and dispersal kernel shapes, and high and low extinction probabilities. In contrast, aggregation increases the steady-state proportion of habitat that is occupied (without climate change). Nonetheless, we find that it is possible to achieve both rapid advance and relatively high patch occupancy when the habitat has a "channeled" pattern, resembling corridors or chains of stepping stones. We adapt techniques from electrical circuit theory to predict the rate of advance efficiently for complex, realistic landscape patterns, whereas the rate cannot be predicted by any simple statistic of aggregation or fragmentation. Conservationists are already advocating corridors and stepping stones as important conservation tools under climate change, but they are vaguely defined and have so far lacked a convincing basis in fundamental population biology. Our work shows how to discriminate properties of a landscape's spatial pattern that affect the speed of colonization (including, but not limited to, patterns like corridors and chains of stepping stones), and properties that affect a species' probability of persistence once established. We can therefore point the way to better land use planning approaches, which will provide functional habitat linkages and also maintain local population viability.

Protected areas facilitate species' range expansions.

The benefits of protected areas (PAs) for biodiversity have been questioned in the context of climate change because PAs are static, whereas the distributions of species are dynamic. Current PAs may, however, continue to be important if they provide suitable locations for species to colonize at their leading-edge range boundaries, thereby enabling spread into new regions. Here, we present an empirical assessment of the role of PAs as targets for colonization during recent range expansions. Records from intensive surveys revealed that seven bird and butterfly species have colonized PAs 4.2 (median) times more frequently than expected from the availability of PAs in the landscapes colonized. Records of an additional 256 invertebrate species with less-intensive surveys supported these findings and showed that 98% of species are disproportionately associated with PAs in newly colonized parts of their ranges. Although colonizing species favor PAs in general, species vary greatly in their reliance on PAs, reflecting differences in the dependence of individual species on particular habitats and other conditions that are available only in PAs. These findings highlight the importance of current PAs for facilitating range expansions and show that a small subset of the landscape receives a high proportion of colonizations by range-expanding species.

Comparing organic farming and land sparing: optimizing yield and butterfly populations at a landscape scale.

Organic farming aims to be wildlife-friendly, but it may not benefit wildlife overall if much greater areas are needed to produce a given quantity of food. We measured the density and species richness of butterflies on organic farms, conventional farms and grassland nature reserves in 16 landscapes. Organic farms supported a higher density of butterflies than conventional farms, but a lower density than reserves. Using our data, we predict the optimal land-use strategy to maintain yield whilst maximizing butterfly abundance under different scenarios. Farming conventionally and sparing land as nature reserves is better for butterflies when the organic yield per hectare falls below 87% of conventional yield. However, if the spared land is simply extra field margins, organic farming is optimal whenever organic yields are over 35% of conventional yields. The optimal balance of land sparing and wildlife-friendly farming to maintain production and biodiversity will differ between landscapes.

Scale matters: the impact of organic farming on biodiversity at different spatial scales.

There is increasing recognition that ecosystems and their services need to be managed in the face of environmental change. However, there is little consensus as to the optimum scale for management. This is particularly acute in the agricultural environment given the level of public investment in agri-environment schemes (AES). Using a novel multiscale hierarchical sampling design, we assess the effect of land use at multiple spatial scales (from location-within-field to regions) on farmland biodiversity. We show that on-farm biodiversity components depend on farming practices (organic vs. conventional) at farm and landscape scales, but this strongly interacts with fine- and coarse-scale variables. Different taxa respond to agricultural practice at different spatial scales and often at multiple spatial scales. Hence, AES need to target multiple spatial scales to maximize effectiveness. Novel policy levers may be needed to encourage multiple land managers within a landscape to adopt schemes that create landscape-level benefits.

Metapopulation responses to patch connectivity and quality are masked by successional habitat dynamics.

Many species have to track changes in the spatial distribution of suitable habitat from generation to generation. Understanding the dynamics of such species will likely require spatially explicit models, and patch-based metapopulation models are potentially appropriate. However, relatively little attention has been paid to developing metapopulation models that include habitat dynamics, and very little to testing the predictions of these models. We tested three predictions from theory about the differences between dynamic habitat metapopulations and their static counterparts using long-term survey data from two metapopulations of the butterfly Plebejus argus. As predicted, we showed first that the metapopulation inhabiting dynamic habitat had a lower level of habitat occupancy, which could not be accounted for by other differences between the metapopulations. Secondly, we found that patch occupancy did not significantly increase with increasing patch connectivity in dynamic habitat, whereas there was a strong positive connectivity-occupancy relationship in static habitat. Thirdly, we found no significant relationship between patch occupancy and patch quality in dynamic habitat, whereas there was a strong, positive quality-occupancy relationship in static habitat. Modeling confirmed that the differences in mean patch occupancy and connectivity-occupancy slope could arise without changing the species' metapopulation parameters-importantly, without changing the dependence of colonization upon connectivity. We found that, for a range of landscape scenarios, successional simulations always produced a lower connectivity-occupancy slope than comparable simulations with static patches, whether compared like-for-like or controlling for mean occupancy. We conclude that landscape-scale studies may often underestimate the importance of connectivity for species occurrence and persistence because habitat turnover can obscure the connectivity-occupancy relationship in commonly available snapshot data.