There is an emerging consensus that many animal species are responding to current climate warming by shifting their distributions northwards. However, in order to track climate, species must be able to disperse through landscapes that have been greatly altered by human activities, and where breeding habitats are often fragmented and scattered across inhospitable urban and agricultural landscapes. Because of this habitat loss, some species are failing to shift and are lagging behind climate changes because they are unable to reach new sites beyond the range margin. This failure to shift is of great concern because many species that are lagging behind climate are endangered species of high conservation concern. Even if greenhouse gas emissions were greatly reduced immediately, more warming would still occur because of inertia in the Earth's climate system. Thus there is a commitment to future warming regardless of any mitigation, and so adaptation measures are required urgently. One commonly suggested adaptation measure is the creation of new habitats as 'corridors' or 'stepping stones' to allow species to move through unsuitable landscapes, and to help them colonise new sites. However, this notion of habitat creation is essentially untested, and there is no comprehensive study of whether such adaptation methods might be successful, or how they might be implemented. The proposed work will provide the first systematic analysis of the importance of habitat availability on distribution lags in an entire group of insects (butterflies). We will develop new dynamic computer models that simulate range expansion through realistic British landscapes. The models take account of the availability of habitat and suitable climate, and also include biological processes such as dispersal, birth and death rates. We will (1) develop and validate dynamic models for all 46 resident, southerly-distributed British butterfly species. We will (2) use these models to quantify the degree to which habitat limits species' range expansion. We will (3) use models to examine the success of different habitat creation scenarios to examine which, if any, scenarios would be sufficient to allow range expansion, and which adaptation strategy is the best. The project will produce results of considerable practical value, as well as addressing fundamental questions about limits to species ranges. It will open up a new avenue of research on the impacts of climate change on biodiversity. Conservation strategies must include adaptation strategies, but conservationists are uncertain about what to do. The proposed work will provide a concrete body of scientific evidence to inform this debate.

Climate change is causing the populations of some species to increase, some to remain relatively stable, and others to decline, even when the species co-exist and might be expected to exhibit comparable ecological responses (e.g., some southern species have expanded their ranges northwards, whereas others have retreated). This diversity of responses to climate change may reflect differences in their capacities to undertake evolutionary and plastic responses that determine success or failure. However, multi-species studies of historical evolutionary responses to environmental change are lacking. In the proposed research, we will use: (1) analyses of historical and present-day DNA from 30 species (10 declining, 10 stable and 10 expanding) to identify the commonality or diversity of adaptive responses to anthropogenic climate change; (2) experimental studies to tease apart plastic, epigenetic and evolutionary responses in a focal species; and (3) modelling to evaluate the contributions of evolutionary, epigenetic and plastic changes to the responses of British Lepidoptera to past and future climatic changes. Moths and butterflies represent an ideal study group because extensive datasets allow us to document the ecological (population abundance, distribution change) and plastic (phenology) responses of species to climate change over the past four decades with a precision not possible for other taxa. Their annual (or faster) generations permit rapid evolutionary change as well as plastic responses to within- and between-year variation in climatic conditions. Museum collections will enable us to assess historical levels of genetic variation within our study species prior to 20th century anthropogenic climate change. We will take advantage of recent advances in sequencing technology to quantify ancestral genetic variation in our study species, and compare this with current genomic diversity to enumerate genetic changes taking place in declining, stable and increasing species, and specifically to evaluate whether species with higher levels of genetic variation show greater ability to adapt to climate change. We will complement this multi-species analysis by evaluating the capacity of expanding, stable and declining populations of one focal species, Pararge aegeria (Speckled wood butterfly) to exhibit evolutionary change, phenotypic plasticity and epigenetic effects using experiments in which we manipulate environmental conditions during larval development (temperature, photoperiod and host-plant desiccation). These experiments will reveal if there are environmental thresholds beyond which adaptive plasticity fails, and the potential for plasticity to evolve and buffer species under future environments. We will then use dynamic simulation models that incorporate our empirical data to test the relative importance of phenotypic plasticity, epigenetic effects, and evolutionary responses in determining species' responses to climate change, and how the relative importance of these factors varies among different species and population types. Once calibrated, we can then use our models to project the responses of these species to future climate change, based on observed limits to adaptation and plasticity. Distinguishing the key factors (ecological, demographic, and genomic) that determine species' responses to environmental change, and how these depend on evolutionary responses, will allow us to identify potential conservation strategies to facilitate population persistence and growth in the face of ongoing climate change.

This proposal falls within the Ecology & Hydrology Funding Initiative. The lead CEH science programme is Biodiversity, but the proposal links strongly to the Environmental Informatics programme and the Climate Change cross-cutting theme There is an emerging consensus that the majority of animal species are responding to climate change, and that many species are at risk of extinction from climate warming. The main metrics of change reported in the scientific literature and in governmental reports are changes in the phenology (the timing of the life cycle, in relation to the changing seasons) and the distributions of species (many British species are moving northwards and to higher elevations). However, these are not the only ways that climate change affects species. The scientific literature and conservation organisations have largely ignored possible climate-driven changes in the habitat associations of species. The temperature conditions experienced by organisms vary between habitats, depending, for example, on the level of shading or protection from frost provided by the vegetation. Therefore, when the climate warms, particular habitats that were too cool in the past (e.g., shady woodlands) may now be warm enough to become occupied, and habitats that used to be favourable may become too hot (e.g., south-facing hillsides). So, the habitats that a species can occupy within a particular region may change as the climate warms and this may affect the patterns of distribution change; there is already evidence that at least a few species are changing their habitat associations in this way. The proposed work will provide the first systematic analysis of the impact of climate change on the habitat associations of an entire group of animals. We will quantify, for 57 species of British butterfly, the degree to which habitat use is modified by climate. We will determine how habitat use varies with geographical variation in the climate, and estimate for the first time how the habitats that species occupy have changed through time (since 1970), as the climate has warmed. The research programme will use the opportunity of climate change to test the ecological hypothesis that species become restricted to particular habitats at the edges of their geographic distributions, where the climate is generally unfavourable. Therefore, we predict that southern species (for which the climate is improving) will have shown increases in the variety of habitats they use over the last 35 years, whereas northern species (for which the climate is deteriorating) will have shown decreases in the range of habitats used. We will also collect new field data to examine relationships between habitat use and species survival in order to investigate the factors influencing species' habitat associations. The project will produce results of considerable practical value. It will open up a new avenue of research on the impacts of climate change on biodiversity. It will provide specific estimates of how the distributions of British butterflies are changing, and therefore whether additional habitats need to be protected, and whether the habitat management that is currently being applied is still appropriate. Conservation strategies must adapt to climate change, but conservationists are uncertain about what to do. The proposed work will provide a concrete body of scientific evidence to inform this debate.

With increasing recognition of the importance of insects, there are growing concerns that insect biodiversity has declined globally, with serious consequences for ecosystem function and services. Yet, gaps in knowledge limit progress in understanding the magnitude and direction of change. Information about insect trends is fragmented, and time-series data are restricted and unrepresentative, both taxonomically and spatially. Moreover, causal links between insect trends and anthropogenic pressures are not well-established. It is, therefore, difficult to evaluate stories about "insectageddon", to understand the ecosystem consequences, to devise mitigation strategies, or predict future trends. To address the shortfalls, we will bring together diverse sources of information, such as meta-analyses, correlative relationships and expert judgement. GLiTRS will collate these diverse lines of evidence on how insect biodiversity has changed in response to anthropogenic pressures, how responses vary according to functional traits, over space, and across biodiversity metrics (e.g. species abundance, occupancy, richness and biomass), and how insect trends drive further changes (e.g. mediated by interaction networks). We will integrate these lines of evidence into a Threat-Response model describing trends in insect biodiversity across the globe. The model will be represented in the form of a series of probabilistic statements (a Bayesian belief network) describing relationships between insect biodiversity and anthropogenic pressures. By challenging this "Threat-Response model" to predict trends for taxa and places where high-quality time series data exist, we will identify insect groups and regions for which indirect data sources are a) sufficient for predicting recent trends, b) inadequate, or c) too uncertain. Knowledge about the predictability of threat-response relationships will allow projections - with uncertainty estimates - of how insect biodiversity has changed globally, across all major taxa, functional groups and biomes. This global perspective on recent trends will provide the basis for an exploration of the consequences of insect decline for a range of ecosystem functions and services, as well as how biodiversity and ecosystem properties might be affected by plausible scenarios of future environmental change. GLiTRS is an ambitious and innovative research program: two features are particularly ground-breaking. First, the collation of multiple forms of evidence will permit a truly global perspective on insect declines that is unachievable using conventional approaches. Second, by validating "prior knowledge" (from evidence synthesis) with recent trends, we will assess the degree to which insect declines are predictable, and at what scales.

Biodiversity represents the life support system on which society depends, but is increasingly threatened by human activities. As a result, there is an urgent need for tested methods to help biodiversity to adapt to emerging threats such as climate change. One possible first step is to protect existing populations of threatened species by making use of the microclimates that are created by different features of the landscape, and that can buffer the effects of climate change. Conservation organisations already carry out activities that influence microclimate, for example by grazing livestock to create hot conditions for plants or invertebrates in short or broken vegetation. Such approaches are based on experience of management actions that have been successful for threatened species until now, but relatively little explicit information is available to guide these management activities under future climate change. We aim to provide practical guidance to help decide what activities should be carried out, where, and for which species, to increase the resilience of biodiversity to climate change We will use techniques developed during recent NERC-funded research, to predict variation in temperature and moisture conditions at a fine resolution throughout the landscape of South West England. We will provide our project partners, the environmental organisations that are charged with conserving biodiversity in the region, with information on how this microclimate variation influences priority species for conservation. We will work closely with these organisations to ensure that the format and content of the resources we develop are practically useful. To achieve this goal, we will liaise closely with the organisations about their conservation priorities in the region, develop a set of microclimate databases to access via Geographic Information Systems, and provide guidance and support to the partners on the application of this resource to conservation planning and management. We will apply the microclimate resources we develop to locations in South-West England in which our partners are guiding conservation management using innovative landscape-scale approaches, which require coordinated management across a range of habitats and land-uses. These landscape-scale projects will provide an opportunity to apply the microclimate information to existing questions of where and how to focus management activities to help protect species against potentially negative effects of climate change. Based on our experience of applying these techniques to conservation management in the landscape-scale projects, we will work with our partners to produce a broader guidance document to assist with planning, prioritisation and management to adapt UK biodiversity conservation to climate change. Our direct beneficiaries are bodies whose primary goal is nature conservation, but we will develop tools and guidance in a format that enables their wider future application by organisations involved with environmental policy, planning and management in the UK.