Understanding within- and between-population variation in responses to climate variability and extreme climatic events

人类活动导致的气候变化导致平均气温上升，这对自然界产生了深远的影响，包括物种相互作用的中断、物种范围的变化、种群数量的减少和物种的灭绝。随着全球变暖，气候变化预计将导致气候变率以及极端气候事件（如热浪）的频率、持续时间和严重程度的变化。与逐渐变暖的气候相比，气候变率的增加和极端事件对生物体的影响可能更为严重，因为它们会导致气候变量（如温度）在短时间尺度上的较大相对变化。在这样做的过程中，他们更有可能将生物体暴露在影响其性能的环境中。因此，我们需要了解气候变率和极端气候事件如何影响生物的生存和繁殖，进化变化为物种适应持续变化提供手段的程度，以及是否存在加剧或减轻气候变率增加或极端气候事件影响的环境因素。了解气候变化和野外极端气候事件影响的进展在很大程度上受到收集适当数据的困难的限制。这是因为这类研究需要与野生种群有关的时空分辨率的气候数据、大量野生个体的生存和繁殖信息、野生系统中个体表型的测量以及资源可用性或栖息地质量等环境特征的详细信息。我们建议使用一个广泛用于研究平均气候变化影响的生态系统-雀形目鸟类在落叶林地中以毛虫为食物来源-来研究气候变率和极端气候事件的变化如何导致生存和繁殖的变化。首先，我们将使用一项对野生动物进行最长时间研究的数据——在牛津附近的威瑟姆森林对大山雀（Parus major）的研究，该研究的数据在标准化条件下存在了近60年。我们将利用这些数据来探索不同空间尺度的气候、栖息地变化的测量之间的联系，以及这些与个体生殖数据和个体适应性之间的联系。然后，我们将通过使用来自欧洲各地27个大山雀研究的数据集来扩展这个单一种群的研究，量化气候变化/极端气候与这些鸟类的生存和繁殖之间关系的全大陆模式，并探索这些关系中种群之间和物种之间的差异。使用上述方法，我们的工作将为以下方面提供新的见解：(1)气候变异和极端气候事件的变化对生存和繁殖的直接影响，从而可能对人口健康产生影响；(2)这些气候变化对当地和欧洲尺度上自然选择的影响；(3)环境异质性对气候变率/极端气候事件影响的调节作用。然后，这些见解可用于预测未来气候变率/极端气候事件的变化如何影响种群和物种，更好地预测进化的潜力，以帮助物种应对持续的人类驱动的气候变化，并查明保护可以利用环境、个体之间和种群之间的变化的方法，以尽量减少持续的气候变化对生物多样性的影响。

Human-driven climate change is leading to increases in average temperature that are having profound impacts on the natural world, including breakdowns in species interactions, shifts in species ranges, and population collapse and species extinctions. Alongside overall warming, climate change is expected to lead to changes in climate variability and in the frequency, duration, and severity of extreme climatic events, such as heat waves. Increased climate variability and extreme events may lead to more severe effects on organisms than more gradual climate warming because they lead to larger relative changes in climatic variables, such as temperature, over short timescales. In doing so, they are more likely to expose organisms to conditions that affect their performance. Consequently, we need to understand how climate variability and extreme climatic events affect the survival and reproduction of organisms, the degree to which evolutionary change provides a means for species to adapt to continued change, and whether there are environmental factors that act to exacerbate or ameliorate the effects of increased climate variability or extreme climatic events. Progress to understand the impacts of climate variability and extreme climatic events in the wild has been limited largely by the difficulty of collecting appropriate data. This is because such studies require climatic data at a temporal and spatial resolution relevant to wild populations, information on the survival and reproduction of large numbers of wild individuals, measures of individual phenotypes in wild systems, and detailed information on environmental characteristics, such as resource availability or habitat quality. We propose to use an ecological system that has been extensively used to study the effects of changing average climate - passerine birds exploiting caterpillars as a food source in deciduous woodlands - to examine how variation in climate variability and extreme climatic events generate variation in survival and reproduction. First, we will use data from one of the longest running studies of a wild animal - the study of great tits (Parus major) in Wytham Woods, near Oxford for which data exist for almost 60 years under standardised conditions. We will use these data to explore the links between climate at a variety of spatial scales, measures of habitat variation, and how these are linked to individual reproductive data and to individual fitness. We will then expand on this single-population study by using datasets from 27 studies of great tits across Europe, to quantify continent-wide patterns in the relationship between climate variability/extremes an survival and reproduction of these birds, and explore differences between populations and between species in these relationships. Using the approach outlined above, our work will provide novel insights into: (1) the immediate consequences of variation in climate variability and extreme climatic events on survival and reproduction, and thus potentially on population health; (2) the influence of these climatic changes on natural selection at both the local and European scale; and (3) the role of environmental heterogeneity in modifying the effects of climate variability/extreme climatic events. These insights can then be used to forecast how future changes in climate variability/extreme climatic events may influence populations and species, better predict the potential for evolution to help species cope with continued human-driven climate change, and pinpoint ways that conservation can use variation in the environment, between individuals, and between populations, to minimise the effects of continued climate change on biodiversity.