ABSTRACT: Small-scale spatial variation in temperature is an important attribute of many habitats, as spatial thermal refugia may allow organisms to avoid some negative consequences of thermal extremes. Although the ecological importance of thermal refugia in intertidal habitats is well-known in summer, the relative availability, effectiveness, and ecological relevance of thermally benign microhabitats in the winter is poorly understood. Here, we explored small-scale thermal variability on a temperate rocky coastline during winter, and the relevance of these thermal patterns to Littorina scutulata, a common intertidal gastropod. Sheltered microhabitats, including crevices, undersides of boulders, and areas beneath algal canopy, were all warmer than surrounding exposed areas during winter low tides. L. scutulata was able to recover relatively quickly from exposure to low temperatures down to -3°C, but performance costs began to accrue at lower temperatures. However, the presence of a refuge habitat buffered the negative effects of cold temperatures, and sheltered L. scutulata regained mobility more quickly than conspecifics on exposed surfaces. Further experiments demonstrated that L. scutulata were more frequently found in thermal refugia when previously exposed to low temperatures, but not when exposed to milder temperatures. Our results suggest that realistic low temperatures can impose important costs on intertidal species such as L. scutulata, but the availability of thermal refugia coupled with behavioural adaptations can allow species to reduce many such costs. Just as in summer, thermal refugia in winter may be important for the distribution and abundance of intertidal species and the diversity and function of coastal ecosystems.

中文翻译：

---

在冬季，低温暴露决定了潮间腹足纲（Littorina scutulata）的性能和热微生境的使用

摘要：温度的小范围空间变化是许多栖息地的重要属性，因为空间热避难所可能使生物体避免极端温度的某些负面影响。尽管在夏季潮间生境中热避难所的生态重要性众所周知，但对冬季热良性微生境的相对可用性，有效性和生态相关性知之甚少。在这里，我们探讨了冬季温带岩石海岸线上的小尺度热变化，以及这些热模式与常见的潮间腹足纲小蜥蜴（Littorina scutulata）的相关性。在冬季退潮期间，包括缝隙，巨石下面以及藻冠下方的区域在内的微生境都比周围的裸露区域温暖。从暴露于低至-3°C的低温下，盾cut菌能够相对快速地恢复，但是在较低的温度下性能成本开始增加。但是，避难所栖息地的存在减轻了低温的负面影响，庇护的盾形乳杆菌比裸露表面上的同种异种更快恢复了活动能力。进一步的实验表明，在先前提到的低温环境中，在热疗中更经常发现盾形乳杆菌，而在较温和的温度下则不常见。我们的结果表明，现实的低温可能会对潮间带的物种（例如s。cutcutata）造成重大损失。，但是热避难所的有效性以及行为适应性可以使物种减少许多此类成本。与夏季一样，冬季的热避风对于潮间带物种的分布和丰富以及沿海生态系统的多样性和功能可能很重要。