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

气候变化正在导致一些物种的种群数量增加，一些物种保持相对稳定，另一些物种数量下降，即使这些物种共存，并可能表现出类似的生态反应(例如，一些南方物种的活动范围向北扩大，而另一些物种已经撤退)。应对气候变化的这种多样性可能反映了它们采取决定成败的进化性和可塑性反应的能力的差异。然而，缺乏对环境变化的历史进化反应的多物种研究。在拟议的研究中，我们将使用：(1)对30个物种(10个衰退物种，10个稳定物种和10个扩张物种)的历史和现在DNA的分析，以确定对人为气候变化的适应反应的共性或多样性；(2)对局部物种的塑料、表观遗传和进化反应进行梳理；以及(3)建立模型，评估进化、表观遗传和塑料变化对英国鳞翅目对过去和未来气候变化的反应的贡献。蛾和蝴蝶是一个理想的研究群体，因为广泛的数据集使我们能够记录过去40年来物种对气候变化的生态(种群丰度、分布变化)和塑料(物候)反应，这是其他类群无法达到的精度。它们的年度(或更快)世代允许快速进化变化，以及对年内和年间气候条件变化的可塑性反应。博物馆的藏品将使我们能够评估在20世纪人为气候变化之前，我们研究的物种中基因变异的历史水平。我们将利用测序技术的最新进展来量化我们研究物种的祖先遗传变异，并将其与当前的基因组多样性进行比较，以列举在衰落、稳定和增长的物种中发生的遗传变化，特别是评估遗传变异水平较高的物种是否表现出更强的适应气候变化的能力。我们将通过评估一个焦点物种Pararge aegeria(斑纹木蝶)的种群扩张、稳定和下降的能力来补充这种多物种分析，通过实验来展示进化变化、表型可塑性和表观遗传效应，在这些实验中，我们操纵幼虫发育期间的环境条件(温度、光周期和寄主植物干燥)。这些实验将揭示是否存在环境阈值，超过了这些阈值，适应性可塑性就会失效，以及可塑性在未来环境下进化和缓冲物种的可能性。然后，我们将使用包含我们的经验数据的动态模拟模型来测试表型可塑性、表观遗传效应和进化反应在确定物种对气候变化的反应中的相对重要性，以及这些因素的相对重要性如何在不同物种和种群类型之间变化。一旦校准，我们就可以使用我们的模型，根据观察到的适应和可塑性限制，预测这些物种对未来气候变化的反应。区分决定物种对环境变化的反应的关键因素(生态、人口和基因组)，以及这些因素如何依赖于进化反应，将使我们能够确定潜在的保护战略，以促进种群在面临持续气候变化时的持续和增长。

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.