Habitat loss and climate change together represent a great threat to biodiversity because species face the difficult task of shifting their distributions across human-dominated landscapes in which suitable habitats are present only as scattered fragments of formerly more widespread types of vegetation. New approaches are needed to understand and predict accurately the responses of species to these environmental drivers of change when they act in combination. The complication is that climate change itself alters habitat quality and quantity, by changing the availability of suitable 'microclimates'. For example, our study species, the silver-spotted skipper butterfly, reaches the cool northern edge of its European distribution in England. As such, it used to be restricted to exceptionally hot microclimates (short grass on South-facing hillsides) in the early 1980s, but has recently colonised cooler habitats (taller grassland, and East, West, and North-facing hillsides) as the climate has warmed. Habitats that used to be too cool are now accessible to the species; although some south-facing grassland may have started to become too hot and dry for the insect. Increasing suitability of East, West, and North-facing hillsides has resulted in a major increase in the amount of grassland that is thermally acceptable, allowing the silver-spotted skipper to start to expand its distribution. However, the situation is complicated because year-to-year variation in climatic conditions constantly alters the suitability of each remaining area of calcareous grassland (depending on the slope, aspect and vegetation). The habitat available to the skipper is a shifting mosaic depending on the weather conditions each year, making it difficult to provide clear guidelines for conservation managers to allow the species to survive and extend its distribution. As the climate changes, this interaction between climate and habitat is likely to complicate the process of conservation planning and habitat management for the many rare species that are now restricted to localised areas of habitat in modern landscapes. To date, the feedback loop between climate and the landscape-scale distribution of habitat has not been incorporated in any scientific modelling framework, but this is required before believable and testable projections of species responses to climate change can be made. We will develop a new approach using a population model that incorporates variation over time in climate-driven habitat availability. These models will be developed using large-scale data on the British distribution, habitat and population sizes of the silver-spotted skipper butterfly for the period 1982 to 2001. The models will then be used to predict post-2001 changes, and we will test our projections against new information on changes in habitat and distribution for the skipper between 2002 and 2010. The project will allow us to test how accurately we can predict changes in species distributions as they respond to climate change, and the importance of climate, habitat and their interactions in explaining the rates at which species extend their distributions. This step is vital to determine whether conservation actions can alleviate the effects of climate change on biodiversity, and which actions are most efficient in this process of adapting conservation to climate change. We will make the software that we develop available to other scientists, policy-makers and conservation practitioners, allowing our approach to be applied to the conservation of the many other rare species facing the same problems as the silver-spotted skipper.

Many species currently survive as localised, refugial populations in regions where they used to be more widespread under more favourable past climatic conditions. These species survive in localised habitats and/or microclimates that are atypical of the surrounding region; for example, a cold-adapted species in Britain might survive on a locally cold, north-facing site when the climate warms. Refugia have been extremely important in allowing species to survive past climatic changes, and are likely to be so again under anthropogenic climate change. Despite this, the local conditions that support population refugia are poorly understood. Thus we have little idea of the attributes (locations, habitats, microclimates) of sites where species may persist in future as the climate changes. Understanding these attributes is vital for informing future conservation policies as well as for developing a deeper fundamental scientific understanding of the dynamics of species' geographic distributions. We will take advantage of the opportunity presented by anthropogenic climate change to observe the creation of refugial populations directly, by studying four species of northerly-distributed butterflies in Britain. Butterflies are ideal study species for this project because there are excellent distribution records in Britain over the past four decades of climate change, and because local microclimate and microhabitat conditions affect all butterfly life stages, from birth to death. We will re-survey sites in Britain for which we have historical distribution data since the 1970s, and which we re-surveyed in 2004-05, to determine where species have survived, and where they have become extinct. We will use dynamic population models that incorporate environmental information for species to identify the local microclimatic and habitat characteristics of locations where populations have survived since the onset of anthropogenic climate change in the 1970s. We will examine the generality of our butterfly findings by studying climate refugial formation in other northern invertebrates. We will then use our models to project the consequences of future climatic changes for species, to the year 2100, and determine the degree to which refugia coincide with the locations of existing protected areas in Britain. The proposed work will provide the first investigation of, and predictive models for, the attributes of locations that promote population persistence in range retreating species. The project will address fundamental questions about the dynamics of species' ranges under climate change, as well as producing results of considerable practical value for policy makers. It will open up a new avenue of research on understanding the impacts of climate change on biodiversity, and provide a concrete body of scientific evidence to inform the debate on developing effective conservation strategies under climate change.

Recent research on the ecological consequences of climate warming has demonstrated that a wide range of vertebrate, invertebrate and plant species have, on average, shifted their distributions towards the poles and to higher elevations. However, individual species vary greatly in their rates of range expansion. The source of this variation is unknown, but is vital to study if we are to understand the fundamental limits on species' distributions, to identify those species that require conservation action, and thereby to devise evidence-based conservation strategies. The proposed research will evaluate why species vary in the rates and directions that their leading-edge (high latitude) range boundaries have shifted over the past 40 years. Our approach will be to focus first on whether climate factors explain most of the observed variation, and then consider habitat factors, using explanatory variables that can be computed and tested across many species and taxa. We will evaluate whether variation in range expansion is determined primarily by variation in the sensitivities of individual species to different components of local climate. We will then address whether suitable habitats are available to colonise, and hence whether this additionally affects species' rates of response. For any remaining unexplained variation, we will then consider dispersal and other species' attributes (body size, development patterns) that might affect the likelihood of species colonising climatically-suitable areas that contain appropriate habitats. We will capitalise on extensive data sets available to examine range changes over the past four decades of climate warming. Our research will primarily use Lepidoptera as our focal model taxon (~ 135 study species). We will test the broader significance of our findings by extending our studies to >300 species from ~14 vertebrate and invertebrate taxa, and by exploring sources of within-species variation in expansion rates (via project PhD studentships). The project has the potential to develop a novel level of fundamental understanding of the dynamics of species' range boundaries and to identify factors constraining species-specific responses to climate change. Outputs from this research will also lead to the development of evidence-based conservation strategies for biodiversity under future climate change.

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 human-induced habitat loss, many species with poor dispersal ability are failing to shift their ranges and are unable to reach new sites beyond their current range margin. Predicting why some species can shift their ranges in response to climate change whilst others cannot, is crucial for improving our projections of species' future distributions. This project will address this issue by investigating species' dispersal behaviour and capability. Even if greenhouse gas emissions were greatly reduced immediately, more warming would still occur due to inertia in the Earth's climate system. Thus, there is a commitment to future warming regardless of any mitigation and, in this context, adaptation measures are required urgently. One commonly suggested adaptation measure is the creation of more permeable landscapes that enable species to movement through degraded landscapes, and help them colonise new sites. However, the effectiveness of improving habitat connectivity for promoting range shifts is essentially untested. There are currently no data examining how species' flight behaviour in response to landscape features may affect their ability to disperse over longer-distances, colonise new sites, and hence shift their ranges. Yet such information will be crucial for understanding the impacts of climate change on the distribution of biodiversity. The proposed work will provide the first investigation of how 'everyday' local flight behaviour in fragmented landscapes translates into longer-distance dispersal and colonisation success. We will focus on butterflies and collect new field data on butterfly flight path characteristics (displacement, speed of flight, directionality, etc) within breeding habitats, within non-breeding habitats, and at habitat/non-habitat patch boundaries. We will incorporate movement information and butterfly behaviour (ovipositing, nectaring, etc) into spatially-explicit dynamic models to estimate movements in 'real' study landscapes. We will test the reliability of our models by comparing modelled movements with those obtained from independent mark-recapture data for the same species and study landscapes. We will then use validated models to examine how variation in flight behaviour and availability of breeding habitat affects the probability of movement in study landscapes. Our models will also allow us to examine the effectiveness of conservation management plans to improve landscape connectivity (Impact Plan). The project will produce results of considerable practical value, as well as addressing fundamental questions about dispersal limits to species ranges. It will open up a new avenue of research on understanding and predicting 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.

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.