The velocity of evolutionary responses of species to ecological change; testing adaptive limits in time and space

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.

Our research will test how biological responses to environmental change depend on the plastic and evolutionary responses of populations to the physical and biological environments they encounter. We will estimate the amount evolutionary change within populations, identify the effects of genetic variation and plasticity on recent evolutionary rates (and vice versa), and evaluate the effects of within-population (genetic variation) and within-individual (plasticity) flexibility in the responses of species to a century of rapid environmental change.

1. Research users and their requirements. The output of our research will have direct relevance to: Government agencies, National NGOs/Conservation Charities, European/Global government and intergovernmental bodies/frameworks, and European/Global NGOs/Conservation Charities.
These organisations are keen to develop conservation strategies that will be robust under climate change. However, they are constrained by a lack of clear scientific guidance on what species, and under what conditions, maintaining genetic and plastic variation within species will be crucial for the persistence of communities and ecosystems.

2. Methods to engage end-users for maximum societal impact.
Partnership Government and NGOs typically require scientific outputs to be re-framed in a manner relevant to conservation policies and actions. We will do this through collaboration between researchers and stakeholders. Methods to engage end-users will include:

a) Co-development of research and publications. We will continue our strategy of co-development and co-authorship of scientific papers with Butterfly Conservation (BC) and using data collected by citizen scientists of the NHM, BC, and NERC CEH.

b) Assessment of need through a working group. We will organise a Knowledge Exchange workshop with ~20 stakeholders, including representatives from Project Partner Butterfly Conservation, as well as SNH, Natural England, JNCC, NRW, Forestry Commission, Defra, Buglife, RSPB, National Trust, National Trust Scotland, Plantlife, BSBI, WWF, UNEP and County Trusts, to present results and co-develop a summary of key results so to achieve maximum impact within their organisations and amongst their memberships.

c) Production of a report to summarise findings according to stakeholder needs. The report will identify factors affecting flexibility in the responses of species to rapid environmental change, and predictions for limits to flexibility in future and identify which types of species are most vulnerable. This will inform appropriate adaptation actions to ensure favourable conservation status of species. The exact form of the report will be shaped at the workshop.

d) Publication and presentation of report to stakeholders.
1. The report: will be published in a form appropriate to stakeholders (summarising major conclusions, with supplementary species-specific online information), the exact form depending on the advice of stakeholders during the workshop.
2. Public discourse: we will present our findings at BC's AGM and give talks to other stakeholders according to advice/invitations received through the workshop.

e) Wider dissemination to the general public. All partners have effective press offices and commitment to dissemination; we have produced six press releases in the last two years, with average success of ~100 press items per release. The NHM and BC will disseminate information through online publications and their communication networks (including links to all other UK and many international museums).