Research in global change ecology relies heavily on global climatic grids derived from estimates of air temperature in open areas at around 2 m above the ground. These climatic grids thus fail to reflect conditions below vegetation canopies and near the ground surface, where critical ecosystem functions are controlled and most terrestrial species reside. Here we provide global maps of soil temperature and bioclimatic variables at a 1-km² resolution for 0-5 and 5-15 cm depth. These maps were created by calculating the difference (i.e., offset) between in-situ soil temperature measurements, based on time series from over 1200 1-km² pixels (summarized from 8500 unique temperature sensors) across all of the world's major terrestrial biomes, and coarse-grained air temperature estimates from ERA5-Land (an atmospheric reanalysis by the European Centre for Medium-Range Weather Forecasts). We show that mean annual soil temperature differs markedly from the corresponding 2 m gridded air temperature, by up to 10°C (mean = 3.0 ± 2.1°C), with substantial variation across biomes and seasons. Over the year, soils in cold and/or dry biomes are substantially warmer (3.6 ± 2.3°C warmer than gridded air temperature), whereas soils in warm and humid environments are on average slightly cooler (0.7 ± 2.3°C cooler). The observed substantial and biome-specific offsets underpin that the projected impacts of climate and climate change on biodiversity and ecosystem functioning are inaccurately assessed when air rather than soil temperature is used, especially in cold environments. The global soil-related bioclimatic variables provided here are an important step forward for any application in ecology and related disciplines. Nevertheless, we highlight the need to fill remaining global gaps by collecting more in-situ measurements of microclimate conditions to further enhance the spatiotemporal resolution of global soil temperature products for ecological applications.

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Research in environmental science relies heavily on global climatic grids derived from estimates of air temperature at around 2 meter above ground1-3. These climatic grids however fail to reflect conditions near and below the soil surface, where critical ecosystem functions such as soil carbon storage are controlled and most biodiversity resides4-8. By using soil temperature time series from over 8500 locations across all of the world’s terrestrial biomes4, we derived global maps of soil temperature-related variables at 1 km resolution for the 0–5 and 5–15 cm depth horizons. Based on these maps, we show that mean annual soil temperature differs markedly from the corresponding 2 m gridded air temperature, by up to 10°C, with substantial variation across biomes and seasons. Soils in cold and/or dry biomes are annually substantially warmer (3.6°C ± 2.3°C) than gridded air temperature, whereas soils in warm and humid environments are slightly cooler (0.7 ± 2.3°C). As a result, annual soil temperature varies less (by 17%) across the globe than air temperature. The effect of macroclimatic conditions on the difference between soil and air temperature highlights the importance of considering that macroclimate warming may not result in the same level of soil temperature warming. Similarly, changes in precipitation could alter the relationship between soil and air temperature, with implications for soil-atmosphere feedbacks9. Our results underpin that the impacts of climate and climate change on biodiversity and ecosystem functioning are inaccurately assessed when air rather than soil temperature is used, especially in cold environments.

Widespread population declines have been reported for diverse Mediterranean butterflies over the last three decades, and have been significantly associated with increased global change impacts. The specific landscape and climatic drivers of these declines remain uncertain for most declining species. Here, we analyse whether plastic phenotypic traits of a model butterfly species (Pieris napi) perform as reliable biomarkers of vulnerability to extreme temperature impacts in natural populations, showing contrasting trends in thermally exposed and thermally buffered populations. We also examine whether improved descriptions of thermal exposure of insect populations can be achieved by combining multiple information sources (i.e., integrating measurements of habitat thermal buffering, habitat thermal amplification, host plant transpiration, and experimental assessments of thermal death time (TDT), thermal avoidance behaviour (TAB) and thermally induced trait plasticity). These integrative analyses are conducted in two demographically declining and two non‐declining populations of P. napi. The results show that plastic phenotypic traits (butterfly body mass and wing size) are reliable biomarkers of population vulnerability to extreme thermal conditions. Butterfly wing size is strongly reduced only in thermally exposed populations during summer drought periods. Laboratory rearing of these populations documented reduced wing size due to significant negative effects of increased temperatures affecting larval growth. We conclude that these thermal biomarkers are indicative of the population vulnerability to increasing global warming impacts, showing contrasting trends in thermally exposed and buffered populations. Thermal effects in host plant microsites significantly differ between populations, with stressful thermal conditions only effectively ameliorated in mid‐elevation populations. In lowland populations, we observe a sixfold reduction in vegetation thermal buffering effects, and larval growth occurs in these populations at significantly higher temperatures. Lowland populations show reduced host plant quality (C/N ratio), reduced leaf transpiration rates and complete above‐ground plant senescence during the peak of summer drought. Amplified host plant temperatures are observed in open microsites, reaching thermal thresholds that can affect larval survival. Overall, our results suggest that butterfly population vulnerability to long‐term drought periods is associated with multiple co‐occurring and interrelated ecological factors, including limited vegetation thermal buffering effects at lowland sites, significant drought impacts on host plant transpiration and amplified leaf surface temperature, as well as reduced leaf quality linked to the seasonal advance of plant phenology. Our results also identify multiannual summer droughts affecting larval growing periods as a key driver of the recently reported butterfly population declines in the Mediterranean biome.

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Ecotones linking open and forested habitats contain multiple microhabitats with varying vegetal structures and microclimatic regimes. Ecotones host many insect species whose development is intimately linked to the microclimatic conditions where they grow (e.g., the leaves of their host plants and the surrounding air). Yet microclimatic heterogeneity at these fine scales and its effects on insects remain poorly quantified for most species. Here we studied how interspecific differences in the use of microhabitats across ecotones lead to contrasting thermal exposure and survival costs between two closely‐related butterflies (Pieris napi and P. rapae). We first assessed whether butterflies selected different microhabitats to oviposit and quantified the thermal conditions at the microhabitat and foliar scales. We also assessed concurrent changes in the quality and availability of host plants. Finally, we quantified larval time of death under different experimental temperatures (thermal death time [TDT] curves) to predict their thermal mortality considering both the intensity and the duration of the microclimatic heat challenges in the field. We identified six processes determining larval thermal exposure at fine scales associated with butterfly oviposition behavior, canopy shading, and heat and water fluxes at the soil and foliar levels. Leaves in open microhabitats could reach temperatures 3–10°C warmer than the surrounding air while more closed microhabitats presented more buffered and homogeneous temperatures. Interspecific differences in microhabitat use matched the TDT curves and the thermal mortality in the field. Open microhabitats posed acute heat challenges that were better withstood by the thermotolerant butterfly, P. rapae, where the species mainly laid their eggs. Despite being more thermosensitive, P. napi was predicted to present higher survivals than P. rapae due to the thermal buffering provided by their selected microhabitats. However, its offspring could be more vulnerable to host‐plant scarcity during summer drought periods. Overall, the different interaction of the butterflies with microclimatic and host‐plant variation emerging at fine scales and their different thermal sensitivity posed them contrasting heat and resource challenges. Our results contribute to setting a new framework that predicts insect vulnerability to climate change based on their thermal sensitivity and the intensity, duration, and accumulation of their heat exposure.

18Vegetation cover generates local microclimatic gradients in the understorey, being especially 19 pronounced at narrow ecotones linking open and forested habitats (open-closed ecotones). 20They provide key habitats for multiple insect communities and may largely determine the 21 exposure of herbivorous insects to the increasing impacts of climate change. We report parallel 22 measurements of microclimatic variables, multiple host-plant traits, and oviposition behaviour 23 in Mediterranean populations of two Pieris butterflies across ecotones of vegetation cover. 24Open microhabitats were significantly warmer, drier, and more exposed to thermal 25 amplification, which increased temperatures to values affecting insect larval survival. Host plants 26 advanced their reproductive phenology and were shorter. Open microhabitats also inhibited the 27 development of shade-adapted plants (e.g. Alliaria petiolata), decreasing fruit production. In 28 contrast, the reproduction of sun-adapted host plants (e.g. Lepidium draba) was vigorous in the 29 open microhabitats and completely inhibited in closed microhabitats, which were exclusively 30 inhabited by non-reproductive ramets. Key plant traits for the selection of oviposition sites by 31 butterflies, such as foliar water and chlorophyll contents, varied significantly across the open-32 closed ecotones. Foliar water content was always lower in the open microhabitats, whereas 33 foliar chlorophyll gradients differed between sun-and shade-adapted plants. The oviposition 34 behavior of Pieris butterflies across the ecotones differed significantly between the 35 thermotolerant species (P. rapae, preferentially selecting open microhabitats) and the 36 thermosensitive species (P. napi, selecting microhabitats protected by vegetation cover), 37 matching the values of thermal susceptibility estimated from parallel heat tolerance assays of 38 the populations. The larvae of the thermotolerant Pieris species grew under completely different 39 thermal conditions due to differential microhabitat selection, indicating marked interspecific 40 differences in thermal exposure (5-10 °C). These results suggest that the impacts of global 41 warming in these communities will likely be mediated by open-closed ecotones, which 42 determine pronounced local variability in thermal exposure, oviposition placement, and host-43 plant traits affecting larval performance in summer. 44 45

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