A major aim of biological research is to understand how new species evolve. Recent research suggests that adaptation to local environmental differences could be the initial stage in the origin of species in diverse groups such as Heliconius butterflies, radiations of island finches, and Lake Victoria cichlid fish. In this model of ecological speciation, natural selection acts on genetic differences that confer improved fitness for a given climate, soil type, local competitor, herbivore or pathogen, and subsequently these locally adapted populations diverge on a trajectory towards being reproductively isolated species. This model could be important in explaining the origin of new parasite species, as parasites will not only have to adapt to the specific habitat, but the range of hosts within a habitat. However the role of ecological speciation in parasites remains largely untested. The objective of this research is to test whether ecological speciation underlies the origin of species in a closely related group of generalist parasitic plants. These species can attach to a range of different host plants with a specialised feeding organ and subsequently extract nutrients and water. We will perform experimental and genetic studies on British native eyebrights (Euphrasia), a groups of 21 parasitic species including taxa of high conservation priority. Our previous research has shown species on isolated islands show extensive hybridisation. This research follows on from this finding, by studying Euphrasia on the isolated island of Fair Isle. This island is a mosaic of different habitats, and as such can be used to test the importance of adaptation to contrasting environments, and whether these parasitic species show preferences for different host plants. We will first test the performance of three ecologically specialised species when grown in different habitats on Fair Isle. To test whether differences in survival reflect adaptation to different hosts, rather than adaptation to other components of the environment, we will grow Euphrasia in an experimental garden site with specific hosts. Finally, we will investigate the genetics of adaptation and speciation by performing genome sequencing of natural populations. This will use high-throughput genomics in conjunction with newer single-molecule sequencing technologies to investigate the genetic changes associated with parasite adaptation. Overall, our results will give important new insights into the origins of parasitic plant species. Our research tackles this issue from multiple angles, revealing the type of natural selection pressure (such as switching host preferences), as well as the genes underlying this variation. More generally, parasitic plants are keystone species of natural systems, as their parasitic nature reduces the vigour of competitively dominant grasses, and thus maintains grassland species diversity. Our work will show whether ecologically specialised Euphrasia are adapted to different hosts, which may in turn affect their use in grassland management.

Human translocation of species and anthropogenic climate change are resulting in some of the fastest rates of species distribution changes ever seen, causing many native and non-native species to be brought together. While the ecological consequences are often well-documented, the evolutionary impacts of hybridization and gene flow between native and non-native species are usually less visible. Yet gene flow between native and non-native species could profoundly affect future evolutionary adaptations and diversification, potentially impacting on species conservation, responses to climate change and the spread of invasive species. Previous studies have only focussed on gene flow between a few exemplar species. So we lack a general understanding of the prevalence and impact of interspecific gene flow across the tree of life, and specifically of how human activities may be altering these rates of gene flow. In this ambitious proposal, we will leverage reference genomes produced by the Darwin Tree of Life project and combine high-throughput sequencing with the latest bioinformatic methods to address a major question of growing importance: What is the extent of gene flow between native and non-native flowering plant species, and is this gene flow of adaptive value to native or non-native species? We will for the first time assess gene flow across a major branch of the tree of life using all 106 native/non-native flowering plant species pairs known to be hybridising in the British Isles. We will then use these data to parameterise models predicting the rate of gene flow between native and non-native species, and test model estimates of cryptic gene flow among species pairs that have not been observed to hybridize. The British flora is intensively studied, and its well characterised distributions, hybrids and ecology make it an ideal model system to build predictive models exploring ecological and genetics factors affecting the rates and effects of gene flow between native and non-native species. Over the course of this project we will generate 24 trillion bases of sequence data, comprising the genomes of 741 individuals from 269 flowering plant species (137 native, 132 non-native). We will use this data to first assess the extent of gene flow between all 106 native/non-native flowering plant species pairs that are known to hybridise in the British Isles (Objective 1), and establish whether the recent range expansions seen in some of these species are associated with increased gene flow (Objective 2). Population samples will then be used to assess evidence for adaptive gene flow in a subset of 10 recipient species that from Objective 1 and 2 show indications of adaptive introgression (Objective 3). We will use the estimates of introgression from Objectives 1 and 2 to build statistical models to understand the genetic and ecological factors affecting gene flow (Objective 4). Finally, we will test assess the accuracy of model predictions of cryptic gene flow between species that are not known to hybridize using additional empirical estimates of gene flow from 17 genera of Asteraceae (daisy family) and Poaceae (grasses) (Objective 5). This project will be a major step towards understanding the evolutionary consequences of human-mediated gene flow between species. Interspecific hybridization could well be widespread, yet gene flow may still be restricted because of postzygotic reproductive barriers. Alternatively, gene flow between species may be common, and with strong fitness consequences. In addition to academic beneficiaries (evolutionary and global change biologists), our results will inform conservation practitioners, control of invasive species, and increase awareness in the general public about the ubiquity and importance of gene flow among species, and evolutionary responses to environmental change.

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.

Climate change represents a challenge to conservation because the species, habitats and other benefits (e.g., soil retention, maintenance of water quality, landscape value) associated with particular nature reserves and other protected areas (e.g. SSSIs) will change. Furthermore, this may undermine the legislative basis of some protected areas that have been designated as important because they support particular species or contain large numbers of individuals of certain species. Government, conservation agencies and volunteers (often through conservation charities) - stakeholders - need to meet this challenge so as to ensure that the limited resources available for conservation are deployed most efficiently. This Knowledge Exchange programme will bring together researchers and stakeholders to identify the questions that stakeholders most require answering to develop conservation strategies that are relevant under climate change, and then to bring together appropriate scientific and other information to answer the key questions identified by the stakeholders. The focus will be on the role of protected areas within conservation strategies. The project will be achieved via networking, workshops and literature / evidence gathering work. The answers will then be disseminated widely through a jointly-produced report, journal article and accounts in stakeholder magazines and web sites; as well as by oral presentations at a launch event and at stakeholder meetings. We will also identify stakeholder requirements for further research and for further Knowledge Exchange activities. The network formed through this programme will be well-placed to drive further integration of science into policy development and conservation action. The network will include researchers at the University of York and NERC Centre for Ecology & Hydrology, Knowledge Exchange specialists, and a variety of stakeholders and policy makers from, e.g., The Royal Society for the Protection of Birds, The Botanical Society of the British Isles, Butterfly Conservation, from the Joint Nature Conservation Committee, Countryside Council for Wales, Natural England and Scottish Natural Heritage, and also from Defra.