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Aim Climatic changes throughout the Pleistocene have strongly modified species distributions. We examine how these range shifts have affected the genetic diversity of a montane butterfly species and whether the genetic diversity in the extant populations is threatened by future climate change. Location Europe. Taxon Erebia epiphron Lepidoptera: Nymphalidae. Methods We analyzed mtDNA to map current genetic diversity and differentiation of E. epiphron across Europe to identify population refugia and postglacial range shifts. We used species distribution modeling (SDM) to hindcast distributions over the last 21,000 years to identify source locations of extant populations and to project distributions into the future (2070) to predict potential losses in genetic diversity. Results We found substantial genetic diversity unique to specific regions within Europe (total number of haplotypes = 31, number of unique haplotypes = 27, H d = 0.9). Genetic data and SDM hindcasting suggest long‐term separation and survival of discrete populations. Particularly, high rates of unique diversity in postglacially colonized sites in England (H d = 0.64) suggest this population was colonized from a now extinct cryptic refugium. Under future climate change, SDMs predict loss of climate suitability for E. epiphron, particularly at lower elevations (, The genetic diversification of cold‐adapted mountain species, such as E. epiphron, has been shaped by Pleistocene glaciations, resulting in unique genetic diversity in isolated populations. The unique genetic diversity in mountain and cold‐adapted species is at under future climate warming, and we predict E. epiphron will lose 38%–64% of its range in the future, resulting in the loss of genetic diversity, reducing its ability to adapt.

Ecological restoration is considered to play an important role in mitigating climate change, protecting biodiversity and preventing environmental degradation. Yet, there are often multiple perspectives on what outcomes restoration should be aiming to achieve, and how we should get to that point. In this paper we interview a range of policy makers, academics and NGO representatives to explore the range of perspectives on the restoration of Indonesia's tropical peatlands – key global ecosystems that have undergone large‐scale degradation. Thematic analysis suggests that participants agreed about the importance of restoration, but had differing opinions on how effective restoration activities to date have been and what a restored peatland landscape should look like. These results exemplify how ecological restoration can mean different things to different people, but also highlight important areas of consensus for moving forward with peatland restoration strategies.

Range shifting is vital for species persistence, but there is little consensus on why individual species vary so greatly in the rates at which their ranges have shifted in response to recent climate warming. Here, using 40 years of distribution data for 291 species from 13 invertebrate taxa in Britain, we show that interactions between habitat availability and exposure to climate change at the range margins explain up to half of the variation in rates of range shift. Habitat generalists expanded faster than more specialised species, but this intrinsic trait explains less of the variation in range shifts than habitat availability, which additionally depends on extrinsic factors that may be rare or widespread at the range margin. Similarly, while climate change likely underlies polewards expansions, we find that more of the between-species variation is explained by differences in habitat availability than by changes in climatic suitability. A model that includes both habitat and climate, and their statistical interaction, explains the most variation in range shifts. We conclude that climate-change vulnerability assessments should focus as much on future habitat availability as on climate sensitivity and exposure, with the expectation that habitat restoration and protection will substantially improve species’ abilities to respond to uncertain future climates.

1. The redistribution of species in response to climate change is expected to significantly challenge environmental management and conservation efforts around the globe. To date, we have had restricted understanding of the benefits and risks that species redistribution may pose to individual countries, and a limited appreciation of the variability in current opportunities for developing effective monitoring approaches that build on existing national frameworks. 2. To assess the present level of ecological, economic and societal risks and opportunities associated with new arrivals of species driven by changes in climatic conditions, we conducted a review of the available information on changes in animal species (both terrestrial and marine) distribution suspected to be linked to climate change in the United Kingdom over the past ten years (2008-2018). 3. We found evidence that at least 55 species have arrived in new locations in the country due to climate change in past decade, with 22 of them suspected to impact positively or negatively the recipient ecosystems, or nearby human communities. Ten of these 55 species were identified using keywords and hashtags on social media. 4. Synthesis and applications. Our work identifies pressing monitoring gaps relevant to the management of species on the move and discusses the potential for social media to help address current information needs. It also calls for more theoretical work to enable the quick identification of species likely to be problematic (or beneficial) and locations likely to experience significant ecological and societal impacts from biodiversity’s redistribution under a changing climate.

Range shifts are a crucial mechanism enabling species to avoid extinction under climate change1,2. The majority of terrestrial biodiversity is concentrated in the tropics3, including species considered most vulnerable to climate warming4, but extensive and ongoing deforestation of tropical forests is likely to impede range shifts5,6. We conduct a global assessment of the potential for tropical species to reach analogous future climates—‘climate connectivity’—and empirically test how this has changed in response to deforestation between 2000 and 2012. We find that over 62% of tropical forest area (~10 million km2) is already incapable of facilitating range shifts to analogous future climates. In just 12 years, continued deforestation has caused a loss of climate connectivity for over 27% of surviving tropical forest, with accelerating declines in connectivity as forest loss increased. On average, if species’ ranges shift as far down climate gradients as permitted by existing forest connectivity, by 2070 they would still experience 0.77 °C of warming under the least severe climate warming scenario and up to 2.6 °C warming for the most severe scenario. Limiting further forest loss and focusing the global restoration agenda towards creating climate corridors are global priorities for improving resilience of tropical forest biotas under climate change. Physically connected habitats are required for terrestrial species to shift their liveable ranges as the tropics warm. The authors show that over half of tropical forest area is currently unable to provide such climate connectivity, and that loss of connectivity is accelerating with deforestation.

This session will critically examine new and emerging approaches to adaptation using three contrasting themes to illustrate key issues. The session will comprise 3 panels on: Conservation/biodiversity, Coastal Systems and Urban Systems. Each panel will address the common themes of (a) co-benefits and transformation issues; (b) how to handle uncertainty; (c) justice dimensions; (d) links between systems; and (e) working across scales – national policy to local implementation.

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.

Temperature is a core component of a species' fundamental niche. At the fine scale over which most organisms experience climate (mm to ha), temperature depends upon the amount of radiation reaching the Earth's surface, which is principally governed by vegetation. Tropical regions have undergone widespread and extreme changes to vegetation, particularly through the degradation and conversion of rainforests. As most terrestrial biodiversity is in the tropics, and many of these species possess narrow thermal limits, it is important to identify local thermal impacts of rainforest degradation and conversion. We collected pantropical, site‐level (

Although the number of studies discerning the impact of climate change on ecological systems continues to increase, there has been relatively little sharing of the lessons learnt when accumulating this evidence. At a recent workshop entitled ‘Using climate data in ecological research’ held at the UK Met Office, ecologists and climate scientists came together to discuss the robust analysis of climate data in ecology. The discussions identified three common pitfalls encountered by ecologists: 1) selection of inappropriate spatial resolutions for analysis; 2) improper use of publically available data or code; and 3) insufficient representation of the uncertainties behind the adopted approach. Here, we discuss how these pitfalls can be avoided, before suggesting ways that both ecology and climate science can move forward. Our main recommendation is that ecologists and climate scientists collaborate more closely, on grant proposals and scientific publications, and informally through online media and workshops. More sharing of data and code (e.g. via online repositories), lessons and guidance would help to reconcile differing approaches to the robust handling of data. We call on ecologists to think critically about which aspects of the climate are relevant to their study system, and to acknowledge and actively explore uncertainty in all types of climate data. And we call on climate scientists to make simple estimates of uncertainty available to the wider research community. Through steps such as these, we will improve our ability to robustly attribute observed ecological changes to climate or other factors, while providing the sort of influential, comprehensive analyses that efforts to mitigate and adapt to climate change so urgently require.

Advances in phenology (the annual timing of species’ life-cycles) in response to climate change are generally viewed as bioindicators of climate change, but have not been considered as predictors of range expansions. Here, we show that phenology advances combine with the number of reproductive cycles per year (voltinism) to shape abundance and distribution trends in 130 species of British Lepidoptera, in response to ~0.5 °C spring-temperature warming between 1995 and 2014. Early adult emergence in warm years resulted in increased within- and between-year population growth for species with multiple reproductive cycles per year (n = 39 multivoltine species). By contrast, early emergence had neutral or negative consequences for species with a single annual reproductive cycle (n = 91 univoltine species), depending on habitat specialisation. We conclude that phenology advances facilitate polewards range expansions in species exhibiting plasticity for both phenology and voltinism, but may inhibit expansion by less flexible species., Many species’ life cycles have moved earlier in the year because of climate change, but we do not know the consequences for range expansions. The authors show that these advances promote range expansions in species with multiple reproductive cycles per year, but not in species with only one.

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., 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.

Both community composition changes due to species redistribution and within-species size shifts may alter body-size structures under climate warming. Here we assess the relative contribution of these processes in community-level body-size changes in tropical moth assemblages that moved uphill during a period of warming. Based on resurvey data for seven assemblages of geometrid moths (>8000 individuals) on Mt. Kinabalu, Borneo, in 1965 and 2007, we show significant wing-length reduction (mean shrinkage of 1.3% per species). Range shifts explain most size restructuring, due to uphill shifts of relatively small species, especially at high elevations. Overall, mean forewing length shrank by ca. 5%, much of which is accounted for by species range boundary shifts (3.9%), followed by within-boundary distribution changes (0.5%), and within-species size shrinkage (0.6%). We conclude that the effects of range shifting predominate, but considering species physiological responses is also important for understanding community size reorganization under climate warming., Body size shifts under climate change may arise from species range shifts, intraspecific size shifts, or both. Here the authors show that body size reduction in moth assemblages on Mt. Kinabalu, Borneo, over 42 years are driven more by species range shifts than by within-species shrinkage.

Many species are extending their leading-edge (cool) range margins polewards in response to recent climate change. In the present study, we investigated range margin changes at the northern (cool) range margins of 1573 southerly-distributed species from 21 animal groups in Great Britain over the past four decades of climate change, updating previous work. Depending on data availability, range margin changes were examined over two time intervals during the past four decades. For four groups (birds, butterflies, macromoths, and dragonflies and damselflies), there were sufficient data available to examine range margin changes over both time intervals. We found that most taxa shifted their northern range margins polewards and this finding was not greatly influenced by changes in recorder effort. The mean northwards range margin change in the first time interval was 23 km per decade (N = 13 taxonomic groups) and, in the second interval, was 18 km per decade (N = 16 taxonomic groups) during periods when the British climate warmed by 0.21 and 0.28 °C per decade, respectively. For the four taxa examined over both intervals, there was evidence for higher rate of range margin change in the more recent time interval in the two Lepidoptera groups. Our analyses confirm a continued range margin shift polewards in a wide range of taxonomic groups.

A cornerstone of conservation is the designation and management of protected areas (PAs): locations often under conservation management containing species of conservation concern, where some development and other detrimental influences are prevented or mitigated. However, the value of PAs for conserving biodiversity in the long term has been questioned given that species are changing their distributions in response to climatic change. There is a concern that PAs may become climatically unsuitable for those species that they were designated to protect, and may not be located appropriately to receive newly-colonizing species for which the climate is improving. In the present study, we analyze fine-scale distribution data from detailed resurveys of seven butterfly and 11 bird species in Great Britain aiming to examine any effect of PA designation in preventing extinctions and promoting colonizations. We found a positive effect of PA designation on species' persistence at trailing-edge warm range margins, although with a decreased magnitude at higher latitudes and altitudes. In addition, colonizations by range expanding species were more likely to occur on PAs even after altitude and latitude were taken into account. PAs will therefore remain an important strategy for conservation. The potential for PA management to mitigate the effects of climatic change for retracting species deserves further investigation.

In our recent Research Article “Individualistic sensitivities and exposure to climate change explain variation in species’ distribution and abundance changes” (1), we presented an analysis relating species-specific measures of sensitivity and exposure to climate, to species’ recent population changes. While our measure and interpretation of species’ climate sensitivity remain correct, we now recognize our interpretation of the exposure measure was inaccurate: Our climate models included an intercept, representing a nonzero average population growth rate; thus, the exposure measure incorporated not only climate effects but other nonclimatic—and potentially unmeasured climatic—effects as well. While our results still demonstrate that a significant proportion of variation in population trends can be explained by exposure and sensitivity, the correct interpretation of the exposure measure means that the explained variation is not solely due to climate. As such, our conclusion that a large proportion of variation in population changes can be explained by individualistic responses to climate is misleading. Given this, and to avoid confusion, we are wholly retracting the Research Article, and we apologize that this was not picked up sooner.

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.

Many species are expanding at their leading-edge range boundaries in response to climate warming. Species are known to respond individualistically to climate change, but there has been little consideration of whether responses are consistent over time. We compared responses of 37 southerly distributed British butterflies over two study periods, first between 1970–1982 and 1995–1999 and then between 1995–1999 and 2005–2009, when mean annual temperature increased regionally by 0.03 °C/yr (a significant rate of increase) and 0.01 °C/yr (a nonsignificant increase) respectively. Our study species might be expected to benefit from climate warming. We measured three responses to climate to investigate this; changes in range margin, distribution area and abundance. In general, the responses of species were inconsistent over time. Species that increased their distribution areas during the first period tended to do so again during the second period, but the relationship was weak. Changes in range margins and abundance were not consistent. In addition, only 5/37 species showed qualitatively similar responses in all three response variables over time (three species increased and two species declined in all variables in both periods). Overall rates of range expansion and distribution area change were significantly greater in the second study period, despite the lower rate of warming, perhaps due to species exploiting climate-distribution lags remaining from the earlier, warmer period. However, there was a significantly greater decline in abundance during the second study period, so range expansions northwards were not necessarily accompanied by increases in distribution area and/or abundance. Hence, species ranges have been thinning as they have expanded northwards. The idiosyncratic responses of these species likely reflect the balance of climatic and habitat drivers of species distribution and abundance changes.

Throughout the world, previously extensive areas of natural habitats have been degraded and fragmented, and improving habitat connectivity may help the long-term persistence of species, and their ability to adapt to climate changes. We focused on Borneo, where many remaining areas of tropical forest are highly fragmented, and we assessed the extent to which Protected Areas (PAs) protect highly-connected forest sites. We analysed remotely-sensed land cover data (0.86 km2 grid cell resolution) using ‘Zonation’ reserve design software, and we ranked grid cells (rank 0–1) according to forest extent and connectivity. PAs currently cover 9% of Borneo, but 60% of Borneo’s total land area lies below 200 m and only 15% of highly-connected cells occurred in these low elevation areas. These findings were relatively insensitive to assumptions about species’ dispersal ability (within the range 1–20 km; representing relatively mobile animal species). The percentage of highly-connected grid cells within PAs could rise from 50% under proposed new PAs (including the ‘Heart of Borneo’ project), although many other highly-connected sites will remain unprotected. On-going land-use changes mean that existing PAs in lowland areas are likely to become increasingly isolated within inhospitable agricultural landscapes, and improving connectivity through reforestation and rehabilitation of degraded forest may be required to maintain the conservation value of these PAs in future.

Creation of new habitat could help species respond to climate change by facilitating range expansion in fragmented landscapes. However, there are currently no guidelines for deciding where new habitat should be placed to promote range changes. We developed a model to simulate the expansion of populations across a heavily fragmented landscape in the United Kingdom, and investigated the effectiveness of six habitat creation strategies for woodland, grassland, heathland, and wetland habitats. A strategy aimed at linking clusters of habitat patches was most effective for three of the four habitat types. Adding habitat evenly or randomly across the landscape, or according to stakeholder suggestions, were consistently better strategies than increasing aggregation of habitat. The results highlight that the best spatial pattern to facilitate range expansion is different from the best pattern to prevent extinction.

During recent climate warming, many insect species have shifted their ranges to higher latitudes and altitudes. These expansions mirror those that occurred after the Last Glacial Maximum when species expanded from their ice age refugia. Postglacial range expansions have resulted in clines in genetic diversity across present-day distributions, with a reduction in genetic diversity observed in a wide range of insect taxa as one moves from the historical distribution core to the current range margin. Evolutionary increases in dispersal at expanding range boundaries are commonly observed in virtually all insects that have been studied, suggesting a positive feedback between range expansion and the evolution of traits that accelerate range expansion. The ubiquity of this phenomenon suggests that it is likely to be an important determinant of range changes. A better understanding of the extent and speed of adaptation will be crucial to the responses of biodiversity and ecosystems to climate change.

Most multicellular terrestrial organisms experience climate at scales of millimetres to metres, yet most species-climate associations are analysed at resolutions of kilometres or more. Because individuals experience heterogeneous microclimates in the landscape, species sometimes survive where the average background climate appears unsuitable, and equally may be eliminated from sites within apparently suitable grid cells where microclimatic extremes are intolerable. Local vegetation structure and topography can be important determinants of fine-resolution microclimate, but a literature search revealed that the vast majority of bioclimate studies do not include fine-scale habitat information, let alone a representation of how habitat affects microclimate. In this paper, we show that habitat type (grassland, heathland, deciduous woodland) is a major modifier of the temperature extremes experienced by organisms. We recorded differences among these habitats of more than 5°C in monthly temperature maxima and minima, and of 10°C in thermal range, on a par with the level of warming expected for extreme future climate change scenarios. Comparable differences were found in relation to variation in local topography (slope and aspect). Hence, we argue that the microclimatic effects of habitat and topography must be included in studies if we are to obtain sufficiently detailed projections of the ecological impacts of climate change to develop detailed adaptation strategies for the conservation of biodiversity.

1.?Current national and international frameworks for assessing threats to species have not been developed in the context of climate change, and are not framed in a way that recognises new opportunities that arise from climate change. 2.?The framework presented here separates the threats and benefits of climate change for individual species. Threat is assessed by the level of climate-related decline within a species’ recently occupied (e.g. pre-1970s) historical distribution, based on observed (e.g. repeat census) and/or projected changes (e.g. modelled bioclimate space). Benefits are assessed in terms of observed and/or projected increases outside the recently occupied historical range. 3.?Exacerbating factors (e.g. small population size, low dispersal capacity) that might increase levels of threat or limit expansion in response to climate change are taken into consideration within the framework. Protocols are also used to identify levels of confidence (and hence research and/or monitoring needs) in each species’ assessment. 4.?Observed and projected changes are combined into single measures of expected decline and increase, together with associated measures of confidence. We weight risk classifications towards information that is most certain. Each species is then placed in one of six categories (high risk, medium risk, limited impact, equivalent risks & benefits, medium benefit, high benefit) reflecting whether climate change is expected (or has been observed) to cause net declines or increases in the region considered, based on the balance of benefits and threats. 5.?We illustrate the feasibility of using the framework by applying it to (i) all British butterflies (N = 58 species) and (ii) an additional sample of British species: 18 species of plants, bats, birds and beetles. 6.?Synthesis. Our framework assesses net declines and increases associated with climate change, for individual species. It could be applied at any scale (regional, continental or global distributions of species), and complements existing conservation assessment protocols such as red-listing. Using observed and projected population and/or range data, it is feasible to carry out systematic conservation status assessments that inform the development of monitoring, adaptation measures and conservation management planning for species that are responding to climate change.

Aim To estimate whether species have shifted at equal rates at their leading edges (cool boundaries) and trailing edges (warm boundaries) in response to climate change. We provide the first such evidence for tropical insects, here examining elevation shifts for the upper and lower boundaries shifts of montane moths. Threats to species on tropical mountains are considered.Location Mount Kinabalu, Sabah, Malaysia.Methods We surveyed Lepidoptera (Geometridae) on Mount Kinabalu in 2007, 42 years after the previous surveys in 1965. Changes in species upper and lower boundaries, elevational extents and range areas were assessed. We randomly subsampled the data to ensure comparable datasets between years. Estimated shifts were compared for endemic versus more widespread species, and for species that reached their range limits at different elevations.Results Species that reached their upper limits at 2500-2700 m (n= 28 species, 20% of those considered) retreated at both their lower and upper boundaries, and hence showed substantial average range contractions (-300 m in elevational extent and -45 km2 in estimated range area). These declines may be associated with changes in cloud cover and the presence of ecological barriers (geological and vegetation transitions) which impede uphill movement. Other than this group, most species (n= 109, 80% of the species considered) expanded their upper boundaries upwards (by an average of 152 m) more than they retreated at their lower boundaries (77 m).Main conclusions Without constraints, leading margins shifted uphill faster than trailing margins retreated, such that many species increased their elevational extents. However, this did not result in increases in range area because the area of land available declines with increasing elevation. Species close to a major ecological/geological transition zone on the mountain flank declined in their range areas. Extinction risk may increase long before species reach the summit, even when undisturbed habitats are available.

Species are thought to have more restricted niches towards their range boundaries, although this has rarely been quantified systematically. We analysed transect data for 41 butterfly species along climatic gradients within Britain and show that 71% of species have broader niches at sites with milder winters. Shifts in habitat associations are considerable across most species' ranges; averaged across all 41 species, we estimate that if 26% of individuals were associated with the favoured habitat on the species' warmest transect, then 70% of individuals would be confined to this habitat on the species' coldest transect. Species with more southerly distributions in Britain showed the greatest changes in their habitat associations. We conclude that geographic variation in realized niche breadth is common and relatively large, especially near range boundaries, and should be taken into account in conserving species under changing climates.

Confidence in projections of the future distributions of species requires demonstration that recently-observed changes could have been predicted adequately. Here we use a dynamic model framework to demonstrate that recently-observed changes at the expanding northern boundaries of three British butterfly species can be predicted with good accuracy. Previous work established that the distributions of the study species currently lag behind climate change, and so we presumed that climate is not currently a major constraint at the northern range margins of our study species. We predicted 1970–2000 distribution changes using a colonisation model, MIGRATE, superimposed on a high-resolution map of habitat availability. Thirty-year rates and patterns of distribution change could be accurately predicted for each species (κ goodness-of-fit of models >0.64 for all three species, corresponding to >83% of grid cells correctly assigned), using a combination of individual species traits, species-specific habitat associations and distance-dependent dispersal. Sensitivity analyses showed that population productivity was the most important determinant of the rate of distribution expansion (variation in dispersal rate was not studied because the species are thought to be similar in dispersal capacity), and that each species’ distribution prior to expansion was critical in determining the spatial pattern of the current distribution. In future, modelling approaches that combine climate suitability and spatially-explicit population models, incorporating demographic variables and habitat availability, are likely to be valuable tools in projecting species’ responses to climatic change and hence in anticipating management to facilitate species’ dispersal and persistence.

Recent climatic change in temperate regions has been rapid and there is mounting speculation that species are failing to keep track of suitable climate, perhaps necessitating assisted colonization for some species. An inability to spread into new areas may result in large reductions in species’ ranges in the future, and threaten the survival of some species. Here we use “speciesclimate” models to predict suitable sites for introductions beyond current range margins, using two U.K. butterfly species. We introduced Melanargia galathea (marbled white) and Thymelicus sylvestris (small skipper) into two sites in northern England, ∼65 and ∼35 km beyond their then-range margins, respectively, to sites that were predicted to be climatically suitable and that appeared to contain suitable habitat for the species. Both introduced populations grew and expanded their range over 6 years (2001–2006; still thriving in 2008), suggesting the existence of a colonization lag and providing evidence that well-planned assisted colonization can be successful. We suggest that assisted colonization may be a feasible and cost-effective means of enabling certain species to track climatic change.

Polewards expansions of species' distributions have been attributed to climate warming, but evidence for climate-driven local extinctions at warm (low latitude/elevation) boundaries is equivocal. We surveyed the four species of butterflies that reach their southern limits in Britain. We visited 421 sites where the species had been recorded previously to determine whether recent extinctions were primarily due to climate or habitat changes. Coenonympha tullia had become extinct at 52% of study sites and all losses were associated with habitat degradation. Aricia artaxerxes was extinct from 50% of sites, with approximately one-third to half of extinctions associated with climate-related factors and the remainder with habitat loss. For Erebia aethiops (extinct from 24% of sites), approximately a quarter of the extinctions were associated with habitat and three-quarters with climate. For Erebia epiphron, extinctions (37% of sites) were attributed mainly to climate with almost no habitat effects. For the three species affected by climate, range boundaries retracted 70–100 km northwards (A. artaxerxes, E. aethiops) and 130–150 m uphill (E. epiphron) in the sample of sites analysed. These shifts are consistent with estimated latitudinal and elevational temperature shifts of 88 km northwards and 98 m uphill over the 19-year study period. These results suggest that the southern/warm range margins of some species are as sensitive to climate change as are northern/cool margins. Our data indicate that climate warming has been of comparable importance to habitat loss in driving local extinctions of northern species over the past few decades; future climate warming is likely to jeopardize the long-term survival of many northern and mountain species.

Evidence is accumulating of shifts in species' distributions during recent climate warming. However, most of this information comes predominantly from studies of a relatively small selection of taxa (i.e., plants, birds and butterflies), which may not be representative of biodiversity as a whole. Using data from less well-studied groups, we show that a wide variety of vertebrate and invertebrate species have moved northwards and uphill in Britain over approximately 25 years, mirroring, and in some cases exceeding, the responses of better-known groups.

Species–climate ‘envelope’ models are widely used to evaluate potential climate change impacts upon species and biodiversity. Previous studies have used a variety of methods to fit models making it difficult to assess relative model performance for different taxonomic groups, life forms or trophic levels. Here we use the same climatic data and modelling approach for 306 European species representing three major taxa (higher plants, insects and birds), and including species of different life form and from four trophic levels. Goodness-of-fit measures showed that useful models were fitted for >96% of species, and that model performance was related neither to major taxonomic group nor to trophic level. These results confirm that such climate envelope models provide the best approach currently available for evaluating reliably the potential impacts of future climate change upon biodiversity.

We investigated the effects of rainfall on the distribution and abundance of the satyrine butterfly Ragadia makuta in selectively logged and unlogged forest on Borneo. In 1997-98, there was a severe El Niño-Southern Oscillation (ENSO) drought, and annual surveys over a 4-y period showed that abundance of R. makuta was greatly reduced during the drought, but that populations quickly recovered after it. Monthly surveys over a 12-mo period of typical rainfall showed that high rainfall in the month preceding surveys significantly reduced butterfly abundance. Butterfly abundance and distribution did not differ between selectively logged and unlogged areas in either monthly or annual surveys and there was no difference between selectively logged and unlogged areas in the pattern of post-drought recovery. These results indicate that the abundance of R. makuta was significantly reduced both after high rainfall and during severe drought, but that these impacts were short-lived and were not affected by habitat disturbance. ENSO droughts on Borneo naturally often lead to widespread forest fires and thus impacts of ENSO events for butterflies are more likely to be due to indirect effects of habitat loss, rather than direct effects of drought on butterfly population dynamics.

New Digs Many studies predict range alterations among species in response to climate change. Species, however, cannot be thought of as truly independent entities, because they all exist as a part of an interacting community. Most discussions of such interactions have focused on the potential restrictions they might place on range expansion; however, Pateman et al. (p. 1028 ) show that climate change has the potential to increase the number of species that can interact and thus facilitate expansion. British data collected by the general public on sightings of the Brown Argus butterfly and its host plants revealed an increased affiliation between the butterflies and a previously little-used group of plants. As summers warmed, populations associated with the new host were more productive than those associated with the more “traditional” host, which facilitated expansion of the butterflies into novel regions.

We analyse distribution records for 51 British butterfly species to investigate altitudinal and latitudinal responses to twentieth century climate warming. Species with northern and/or montane distributions have disappeared from low elevation sites and colonized sites at higher elevations during the twentieth century, consistent with a climate explanation. We found no evidence for a systematic shift northwards across all species, even though 11 out of 46 southerly distributed species have expanded in the northern part of their distributions. For a subset of 35 species, we model the role of climate in limiting current European distributions and predict potential future distributions for the period 2070-2099. Most northerly distributed species will have little opportunity to expand northwards and will disappear from areas in the south, resulting in reduced range sizes. Southerly distributed species will have the potential to shift northwards, resulting in similar or increased range sizes. However, 30 out of 35 study species have failed to track recent climate changes because of lack of suitable habitat, so we revised our estimates accordingly for these species and predicted 65% and 24% declines in range sizes for northern and southern species, respectively. These revised estimates are likely to be more realistic predictions of future butterfly range sizes.

Habitat degradation and climate change are thought to be altering the distributions and abundances of animals and plants throughout the world, but their combined impacts have not been assessed for any species assemblage1,2,3,4. Here we evaluated changes in the distribution sizes and abundances of 46 species of butterflies that approach their northern climatic range margins in Britain—where changes in climate and habitat are opposing forces. These insects might be expected to have responded positively to climate warming over the past 30 years, yet three-quarters of them declined: negative responses to habitat loss have outweighed positive responses to climate warming. Half of the species that were mobile and habitat generalists increased their distribution sites over this period (consistent with a climate explanation), whereas the other generalists and 89% of the habitat specialists declined in distribution size (consistent with habitat limitation). Changes in population abundances closely matched changes in distributions. The dual forces of habitat modification and climate change are likely to cause specialists to decline, leaving biological communities with reduced numbers of species and dominated by mobile and widespread habitat generalists.

Climate warming threatens the survival of species at their warm, trailing-edge range boundaries but also provides opportunities for the ecological release of populations at the cool, leading edges of their distributions. Thus, as the climate warms, leading-edge populations are expected to utilize an increased range of habitat types, leading to larger population sizes and range expansion. Here, we test the hypothesis that the habitat associations of British butterflies have expanded over three decades of climate warming. We characterize the habitat breadth of 27 southerly distributed species from 77 monitoring transects between 1977 and 2007 by considering changes in densities of butterflies across 11 habitat types. Contrary to expectation, we find that 20 of 27 (74%) butterfly species showed long-term contractions in their habitat associations, despite some short-term expansions in habitat breadth in warmer-than-usual years. Thus, we conclude that climatic warming has ameliorated habitat contractions caused by other environmental drivers to some extent, but that habitat degradation continues to be a major driver of reductions in habitat breadth and population density of butterflies.

The distributions of many terrestrial organisms are currently shifting in latitude or elevation in response to changing climate. Using a meta-analysis, we estimated that the distributions of species have recently shifted to higher elevations at a median rate of 11.0 meters per decade, and to higher latitudes at a median rate of 16.9 kilometers per decade. These rates are approximately two and three times faster than previously reported. The distances moved by species are greatest in studies showing the highest levels of warming, with average latitudinal shifts being generally sufficient to track temperature changes. However, individual species vary greatly in their rates of change, suggesting that the range shift of each species depends on multiple internal species traits and external drivers of change. Rapid average shifts derive from a wide diversity of responses by individual species.

Species-energy theory indicates that recent climate warming should have driven increases in species richness in cool and species-poor parts of the Northern Hemisphere. We confirm that the average species richness of British butterflies has increased since 1970–82, but much more slowly than predicted from changes of climate: on average, only one-third of the predicted increase has taken place. The resultant species assemblages are increasingly dominated by generalist species that were able to respond quickly. The time lag is confirmed by the successful introduction of many species to climatically suitable areas beyond their ranges. Our results imply that it may be decades or centuries before the species richness and composition of biological communities adjusts to the current climate.

During recent climate warming, some species have expanded their ranges northwards to keep track of climate changes. Evolutionary changes in dispersal have been demonstrated in these expanding populations and here we show that increased dispersal is associated with reduced investment in reproduction in populations of the speckled wood butterfly, Pararge aegeria. Evolutionary changes in flight versus reproduction will affect the pattern and rate of expansion at range boundaries in the future, and understanding these responses will be crucial for predicting the distribution of species in the future as climates continue to warm.

In order to make realistic predictions of species’ responses to future climate change we need to understand the relative importance of biotic versus abiotic factors in limiting species distributions. We focus on British butterflies, a group of species for which there are good current and historical distribution records. We review our previous studies investigating the relative importance of climate and habitat availability in limiting butterfly distributions. Our studies have used a combination of modelling and analysis of distribution records to investigate factors determining limits to species’ distributions in Europe, and to investigate recent range expansions of butterflies in Britain. Climates in Europe have warmed during the 20th century and many northern areas are improving for butterflies in terms of climate suitability. However, the widespread loss and fragmentation of natural habitats means that many climatically suitable areas are beyond the reach of dispersing adults and so species are unable to keep track of climate changes. In the future, many species may have the potential to occupy many northerly regions that are currently unsuitable. However, most of these newly available areas are remote from current distributions and many species are unlikely to be able to keep track of rapidly warming future climates.

Evidence of anthropogenic global climate change is accumulating, but its potential consequences for insect distributions have received little attention. We use a 'climate response surface' model to investigate distribution changes at the northern margin of the speckled wood butterfly, Pararge aegeria. We relate its current European distribution to a combination of three bioclimatic variables. We document that P. aegeria has expanded its northern margin substantially since 1940, that changes in this species' distribution over the past 100 years are likely to have been due to climate change, and that P. aegeria will have the potential to shift its range margin substantially northwards under predicted future climate change. At current rates of expansion, this species could potentially colonize all newly available climatically suitable habitat in the UK over the next 50 years or more. However, fragmentation of habitats can affect colonization, and we show that availability of habitat may be constraining range expansion of this species at its northern margin in the UK. These lag effects may be even more pronounced in less-mobile species inhabiting more fragmented landscapes, and highlight how habitat distribution will be crucial in predicting species' responses to future climate change.

Until recently, published evidence for the responses of species to climate change had revealed more examples of species expanding than retracting their distributions. However, recent papers on butterflies and frogs now show that population-level and species-level extinctions are occurring. The relative lack of previous information about range retractions and extinctions appears to stem, at least partly, from a failure to survey the distributions of species at sufficiently fine resolution to detect declines, and from a failure to attribute such declines to climate change. The new evidence suggests that climate-driven extinctions and range retractions are already widespread.

Analysis of butterfly and moth species shows that responses to climate change may be more predictable than previously recognized., The responses of animals and plants to recent climate change vary greatly from species to species, but attempts to understand this variation have met with limited success. This has led to concerns that predictions of responses are inherently uncertain because of the complexity of interacting drivers and biotic interactions. However, we show for an exemplar group of 155 Lepidoptera species that about 60% of the variation among species in their abundance trends over the past four decades can be explained by species-specific exposure and sensitivity to climate change. Distribution changes were less well predicted, but nonetheless, up to 53% of the variation was explained. We found that species vary in their overall sensitivity to climate and respond to different components of the climate despite ostensibly experiencing the same climate changes. Hence, species have undergone different levels of population “forcing” (exposure), driving variation among species in their national-scale abundance and distribution trends. We conclude that variation in species’ responses to recent climate change may be more predictable than previously recognized.

Funder: Royal Entomological Society; Id: http://dx.doi.org/10.13039/100013849, Funder: Rural & Environment Science & Analytical Services Division of the Scottish Government, Funder: Alexander von Humboldt Foundation; Id: http://dx.doi.org/10.13039/100005156, Funder: Department of Agriculture, Environment & Rural Affairs, Funder: Center for Environmental and Sustainability Research, 1. Entomology is key to understanding terrestrial and freshwater ecosystems at a time of unprecedented anthropogenic environmental change, and offers substantial untapped potential to benefit humanity in a variety of ways, from improving agricultural practices to managing vector-borne diseases and inspiring technological advances. 2. We identified high priority challenges for entomology using an inclusive, open, and democratic four-stage prioritisation approach, conducted among the membership and affiliates (hereafter ‘members’) of the UK-based Royal Entomological Society (RES). 3. A list of 710 challenges was gathered from 189 RES members. Thematic analysis was used to group suggestions, followed by an online vote to determine initial priorities, which were subsequently ranked during an online workshop involving 37 participants. 4. The outcome was a set of 61 priority challenges within four groupings of related themes: (i) ‘Fundamental Research’ (themes: Taxonomy, ‘Blue Skies’ [defined as research ideas without immediate practical application], Methods and Techniques); (ii) ‘Anthropogenic Impacts and Conservation’ (themes: Anthropogenic Impacts, Conservation Options); (iii) ‘Uses, Ecosystem Services and Disservices’ (themes: Ecosystem Benefits, Technology and Resources [use of insects as a resource, or as inspiration], Pests); (iv) ‘Collaboration, Engagement and Training’ (themes: Knowledge Access, Training and Collaboration, Societal Engagement). 5. Priority challenges encompass research questions, funding objectives, new technologies, and priorities for outreach and engagement. Examples include training taxonomists, establishing a global network of insect monitoring sites, understanding the extent of insect declines, exploring roles of cultivated insects in food supply chains, and connecting professional with amateur entomologists. Responses to different challenges could be led by amateur and professional entomologists, at all career stages. 6. Overall, the challenges provide a diverse array of options to inspire and initiate entomological activities, and reveal the potential of entomology to contribute to addressing global challenges related to human health and well-being, and environmental change.

There is little consensus as to why there is so much variation in the rates at which different species’ geographic ranges expand in response to climate warming[1,2]. Here, we show for British butterfly species that the relative importance of species’ abundance trends and habitat availability vary over time. Species with high habitat availability expanded more rapidly from the 1970s to mid-1990s, when abundances were generally stable, whereas habitat availability effects were confined to the subset of species with stable abundances from the mid-1990s to 2009, when abundance trends were generally declining. This suggests that stable (or positive) abundance trends are a prerequisite for range expansion. Given that species’ abundance trends vary over time[3] for non-climatic as well as climatic reasons, assessment of abundance trends will help improve predictions of species’ responses to climate change, and help understand the likely success of different conservation strategies for facilitating their expansions.