• Energy Research

Summary Species are altering their phenology to track warming temperatures. In forests, understorey plants experience tree canopy shading resulting in light and temperature conditions, which strongly deviate from open habitats. Yet, little is known about understorey phenology responses to forest microclimates. We recorded flowering onset, peak, end and duration of 10 temperate forest understorey plant species in two mesocosm experiments to understand how phenology is affected by sub‐canopy warming and how this response is modulated by illumination, which is related to canopy change. Furthermore, we investigated whether phenological sensitivities can be explained by species' characteristics, such as thermal niche. We found a mean advance of flowering onset of 7.1 d per 1°C warming, more than previously reported in studies not accounting for microclimatic buffering. Warm‐adapted species exhibited greater advances. Temperature sensitivity did not differ between early‐ and later‐flowering species. Experimental illumination did not significantly affect species' phenological temperature sensitivities, but slightly delayed flowering phenology independent from warming. Our study suggests that integrating sub‐canopy temperature and light availability will help us better understand future understorey phenology responses. Climate warming together with intensifying canopy disturbances will continue to drive phenological shifts and potentially disrupt understorey communities, thereby affecting forest biodiversity and functioning.

Summary Species are altering their phenology to track warming temperatures. In forests, understorey plants experience tree canopy shading resulting in light and temperature conditions, which strongly deviate from open habitats. Yet, little is known about understorey phenology responses to forest microclimates. We recorded flowering onset, peak, end and duration of 10 temperate forest understorey plant species in two mesocosm experiments to understand how phenology is affected by sub‐canopy warming and how this response is modulated by illumination, which is related to canopy change. Furthermore, we investigated whether phenological sensitivities can be explained by species' characteristics, such as thermal niche. We found a mean advance of flowering onset of 7.1 d per 1°C warming, more than previously reported in studies not accounting for microclimatic buffering. Warm‐adapted species exhibited greater advances. Temperature sensitivity did not differ between early‐ and later‐flowering species. Experimental illumination did not significantly affect species' phenological temperature sensitivities, but slightly delayed flowering phenology independent from warming. Our study suggests that integrating sub‐canopy temperature and light availability will help us better understand future understorey phenology responses. Climate warming together with intensifying canopy disturbances will continue to drive phenological shifts and potentially disrupt understorey communities, thereby affecting forest biodiversity and functioning.

Forest microclimates contrast strongly with the climate outside forests. To fully understand and better predict how forests' biodiversity and functions relate to climate and climate change, microclimates need to be integrated into ecological research. Despite the potentially broad impact of microclimates on the response of forest ecosystems to global change, our understanding of how microclimates within and below tree canopies modulate biotic responses to global change at the species, community and ecosystem level is still limited. Here, we review how spatial and temporal variation in forest microclimates result from an interplay of forest features, local water balance, topography and landscape composition. We first stress and exemplify the importance of considering forest microclimates to understand variation in biodiversity and ecosystem functions across forest landscapes. Next, we explain how macroclimate warming (of the free atmosphere) can affect microclimates, and vice versa, via interactions with land-use changes across different biomes. Finally, we perform a priority ranking of future research avenues at the interface of microclimate ecology and global change biology, with a specific focus on three key themes: (1) disentangling the abiotic and biotic drivers and feedbacks of forest microclimates; (2) global and regional mapping and predictions of forest microclimates; and (3) the impacts of microclimate on forest biodiversity and ecosystem functioning in the face of climate change. The availability of microclimatic data will significantly increase in the coming decades, characterizing climate variability at unprecedented spatial and temporal scales relevant to biological processes in forests. This will revolutionize our understanding of the dynamics, drivers and implications of forest microclimates on biodiversity and ecological functions, and the impacts of global changes. In order to support the sustainable use of forests and to secure their biodiversity and ecosystem services for future generations, microclimates cannot be ignored. Peer reviewed

Forest microclimates contrast strongly with the climate outside forests. To fully understand and better predict how forests' biodiversity and functions relate to climate and climate change, microclimates need to be integrated into ecological research. Despite the potentially broad impact of microclimates on the response of forest ecosystems to global change, our understanding of how microclimates within and below tree canopies modulate biotic responses to global change at the species, community and ecosystem level is still limited. Here, we review how spatial and temporal variation in forest microclimates result from an interplay of forest features, local water balance, topography and landscape composition. We first stress and exemplify the importance of considering forest microclimates to understand variation in biodiversity and ecosystem functions across forest landscapes. Next, we explain how macroclimate warming (of the free atmosphere) can affect microclimates, and vice versa, via interactions with land-use changes across different biomes. Finally, we perform a priority ranking of future research avenues at the interface of microclimate ecology and global change biology, with a specific focus on three key themes: (1) disentangling the abiotic and biotic drivers and feedbacks of forest microclimates; (2) global and regional mapping and predictions of forest microclimates; and (3) the impacts of microclimate on forest biodiversity and ecosystem functioning in the face of climate change. The availability of microclimatic data will significantly increase in the coming decades, characterizing climate variability at unprecedented spatial and temporal scales relevant to biological processes in forests. This will revolutionize our understanding of the dynamics, drivers and implications of forest microclimates on biodiversity and ecological functions, and the impacts of global changes. In order to support the sustainable use of forests and to secure their biodiversity and ecosystem services for future generations, microclimates cannot be ignored. Peer reviewed

Abstract Global forest cover is heavily fragmented. Due to high edge-to-surface ratios in small forest patches, a large proportion of forests is affected by edge influences involving steep microclimatic gradients. Although forest edges are important ecotones and account for 20% of the global forested area, it remains unclear how biotic and abiotic drivers affect forest edge microclimates at the continental scale. Here we report soil and air temperatures measured in 225 deciduous forest plots across Europe for two years. Forest stands were situated along a latitudinal gradient and subject to a varying vegetation structure as quantified by terrestrial laser scanning. In summer, the average offset of air and soil temperatures in forest edges compared to temperatures outside the forest amounted to -2.8 °C and -2.3 °C, respectively. Edge-to-interior summer temperature gradients were affected by the macroclimate and edge structure. From the edge onwards, larger offsets were observed in dense forest edges and in warmer, southern regions. In open forests and northern Europe, altered microclimatic conditions extended deeper into the forest and gradients were steeper. Canopy closure and plant area index were important drivers of summer offsets in edges, whereas in winter also the forest-floor biomass played a key role. Using high-resolution maps, we estimated that approximately 10% of the European broadleaved forests would be affected by altered temperature regimes. Gradual transition zones between forest and adjacent lands are valuable habitat types for edge species. However, if cool and moist forest interiors are desired, then (i) dense and complex forest edges, (ii) an undisturbed forested buffer zone of at least 12.5 m deep and (iii) trees with a high shade casting ability could all contribute to an increased offset. These findings provide important guidelines to mitigate edge influences, to protect typical forest microclimates and to adapt forest management to climate change.

Abstract Global forest cover is heavily fragmented. Due to high edge-to-surface ratios in small forest patches, a large proportion of forests is affected by edge influences involving steep microclimatic gradients. Although forest edges are important ecotones and account for 20% of the global forested area, it remains unclear how biotic and abiotic drivers affect forest edge microclimates at the continental scale. Here we report soil and air temperatures measured in 225 deciduous forest plots across Europe for two years. Forest stands were situated along a latitudinal gradient and subject to a varying vegetation structure as quantified by terrestrial laser scanning. In summer, the average offset of air and soil temperatures in forest edges compared to temperatures outside the forest amounted to -2.8 °C and -2.3 °C, respectively. Edge-to-interior summer temperature gradients were affected by the macroclimate and edge structure. From the edge onwards, larger offsets were observed in dense forest edges and in warmer, southern regions. In open forests and northern Europe, altered microclimatic conditions extended deeper into the forest and gradients were steeper. Canopy closure and plant area index were important drivers of summer offsets in edges, whereas in winter also the forest-floor biomass played a key role. Using high-resolution maps, we estimated that approximately 10% of the European broadleaved forests would be affected by altered temperature regimes. Gradual transition zones between forest and adjacent lands are valuable habitat types for edge species. However, if cool and moist forest interiors are desired, then (i) dense and complex forest edges, (ii) an undisturbed forested buffer zone of at least 12.5 m deep and (iii) trees with a high shade casting ability could all contribute to an increased offset. These findings provide important guidelines to mitigate edge influences, to protect typical forest microclimates and to adapt forest management to climate change.

Abstract Most biodiversity dynamics and ecosystem processes on land take place in microclimates that are decoupled from the climate as measured by standardised weather stations in open, unshaded locations. As a result, microclimate monitoring is increasingly being integrated in many studies in ecology and evolution. Overviews of the protocols and measurement methods related to microclimate are needed, especially for those starting in the field and to achieve more generality and standardisation in microclimate studies. Here, we present 10 practical guidelines for ground‐based research of terrestrial microclimates, covering methods and best practices from initial conceptualisation of the study to data analyses. Our guidelines encompass the significance of microclimates; the specifics of what, where, when and how to measure them; the design of microclimate studies; and the optimal approaches for analysing and sharing data for future use and collaborations. The paper is structured as a chronological guide, leading the reader through each step necessary to conduct a comprehensive microclimate study. At the end, we also discuss further research avenues and development in this field. With these 10 guidelines for microclimate monitoring, we hope to stimulate and advance microclimate research in ecology and evolution, especially under the pressing need to account for buffering or amplifying abilities of contrasting microhabitats in the context of global climate change.