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Agricultural areas represent one of the major ecosystems of the world. Intensification of agricultural practices produced openfields characterized by low biological diversity. Nevertheless, the distance up to which intensive agricultural fields alter surrounding natural systems is rarely quantified. We determined the spatial scale at which agricultural landscapes alter the diversity of Odonates, a key taxon in wetland ponds, and we tested to what extent citizen science data can be used reliably for this purpose. We compiled 7731 observations made in a portion of the region Centre-Val-de-Loire (France) over 10 years by naturalists on 729 water bodies to analyze the effect of agricultural landscapes (mainly wheat, rapeseed, sunflower) on the species richness of both damselflies and dragonflies in lentic systems. Sixty species were reported over the 10-year period. For dragonflies, intensive agricultural landscapes best explained their richness at the scales of 800 and 1600 m for overall and autochthonous species, respectively, when using the full dataset. The spatial scale was smaller for damselflies, at 200 m for both overall and autochthonous species. These distances were not severely impacted when constraining the data to consider several biases. Multimodel averaging showed that the proportion of intensive agriculture decreased species richness, despite the potential biases inherent to an imperfect database acquired by citizens. This imperfect citizen dataset allows to infer the lowest effect size of agriculture on species richness. Quantitatively, this effect was more important for autochthonous species. Interestingly, both relatively rare taxa and common or generalist species can be under threat in intensive agricultural landscapes, calling for more ecotoxicological studies. The influence of agricultural practices from a distance implies that conservation and management plans of wetland ponds should consider the landscape ecological characteristics and not only the pond features. Conservation efforts focusing too locally on a site may be undermined because intensive agriculture from a distance limits the potential for the site to recover highly diverse communities. These distant effects should be integrated by policy-makers when deciding which wetland pond should benefit from a conservation plan or which conservation action may be planned, implementing, for instance, buffer zones and/or ecological corridors composed of natural vegetation., (© 2024 The Author(s). Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Thermal limits are often used as proxies to assess the vulnerability of ectotherms to environmental change. While meta-analyses point out a relatively low plasticity of heat limits and a large interspecific variability, only few studies have compared the heat tolerance of interacting species. The present study focuses on the thermal limits, and their plasticity (heat hardening), of three species co-occurring in Western Africa: two ectoparasitoid species, Dinarmus basalis (Rondani) (Hymenoptera: Pteromalidae) and Eupelmus vuilleti (Crawford) (Hymenoptera: Eupelmidae), and their common host, Callosobruchus maculatus (F.) (Coleoptera: Bruchidae). The investigation delves into the Critical Thermal Maximum (CTmax), representing the upper tolerance limit, to understand how these species may cope with extreme thermal events. The CTmax of all three species appeared similarly high, hovering around 46.5 °C, exceeding the global mean CTmax observed in insects by 3.5 °C. Short-term exposure to moderate heat stress showed no impact on CTmax, suggesting a potential lack of heat hardening in these species. Therefore, we emphasized the similarity of heat tolerance in these interacting species, potentially stemming from both evolutionary adaptations to high temperatures during development and the stable and similar microclimate experienced by the three species over the years. While the high thermal tolerance should allow these species to endure extreme temperature events, the apparent lack of plasticity raises concerns about their ability to adapt to future climate change scenarios. Overall, this research provides valuable insights into the thermal physiology of these interacting species, providing a basis for understanding their responses to climate change and potential implications for the host-parasitoid system., Competing Interests: Declaration of competing interest None., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Phenological models for insect pests often rely on knowledge of thermal reaction norms. These may differ in shape depending on developmental thermal conditions (e.g. constant vs. fluctuating) and other factors such as life-stages. Here, we conducted an extensive comparative study of the thermal reaction norms for development and viability in the invasive fly, Drosophila suzukii, under constant and fluctuating thermal regimes. Flies, were submitted to 15 different constant temperatures (CT) ranging from 8 to 35 °C. We compared responses under CT with patterns observed under 15 different fluctuating temperature (FT) regimes. We tested several equations for thermal performance curves and compared various models to obtain thermal limits and degree-day estimations. To validate the model's predictions, the phenology was monitored in two artificial field-like conditions and two natural conditions in outdoor cages during spring and winter. Thermal reaction norm for viability from egg to pupa was broader than that from egg to adult. FT conditions yielded a broader thermal breadth for viability than CT, with a performance extended towards the colder side, consistent with our field observations in winter. Models resulting from both CT and FT conditions made accurate predictions of degree-day as long as the temperature remained within the linear part of the developmental rate curve. Under cold artificial and natural winter conditions, a model based on FT data made more accurate predictions. Model based on CT failed to predict adult's emergence in winter. We also document the first record of development and adult emergence throughout winter in D. suzukii. Population dynamics models in D. suzukii are all based on summer phenotype and CT. Accounting for variations between seasonal phenotypes, stages, and thermal conditions (CT vs. FT) could improve the predictive power of the models., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Most ectotherms rely on behavioural thermoregulation to maintain body temperatures close to their physiological optimum. Hence, ectotherms can drastically limit their exposure to thermal extremes by selecting a narrower range of temperatures, which includes their preferred temperature (Tpref). Despite evidence that behavioural thermoregulation can be adjusted by phenotypic plasticity or constrained by natural selection, intraspecific Tpref variations across environmental gradients remain overlooked as compared to other thermal traits like thermal tolerance. Here, we analyzed Tpref variation of spider populations found along a gradient of urban heat island (UHI) which displays large thermal variations over small distances. We measured two components of the thermal preference, namely the mean Tpref and the Tpref range (i.e., standard deviation) in 557 field-collected individuals of a common ground-dwelling spider (Pardosa saltans, Lycosidae) using a laboratory thermal gradient. We determined if Tpref values differed among ten populations from contrasting thermal zones. We showed that endogenous factors such as body size or sex primarily determine both mean Tpref and Tpref range. The Tpref range was also linked to the UHI intensity to a lesser extent, yet only in juveniles. The absence of relationship between Tpref metrics and UHI in adult spiders suggests a Bogert effect according to which the ability of individuals to detect and exploit optimal microclimates weakens the selection pressure of temperatures (here driven by UHI) on their thermal physiology. Alternatively, this lack of relationship could also indicate that temperature patterns occurring at the scale of the spiders' micro-habitat differ from measured ones. This study shows the importance of considering both inter-individual and inter-population variations of the Tpref range when conducting Tpref experiments, and supports Tpref range as being a relevant measure to inform on the strength of behavioural thermoregulation in a given population., Competing Interests: Declaration of competing interest We declare no competing financial interest., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

A core challenge in global change biology is to predict how species will respond to future environmental change and to manage these responses. To make such predictions and management actions robust to novel futures, we need to accurately characterize how organisms experience their environments and the biological mechanisms by which they respond. All organisms are thermodynamically connected to their environments through the exchange of heat and water at fine spatial and temporal scales and this exchange can be captured with biophysical models. Although mechanistic models based on biophysical ecology have a long history of development and application, their use in global change biology remains limited despite their enormous promise and increasingly accessible software. We contend that greater understanding and training in the theory and methods of biophysical ecology is vital to expand their application. Our review shows how biophysical models can be implemented to understand and predict climate change impacts on species' behavior, phenology, survival, distribution, and abundance. It also illustrates the types of outputs that can be generated, and the data inputs required for different implementations. Examples range from simple calculations of body temperature at a particular site and time, to more complex analyses of species' distribution limits based on projected energy and water balances, accounting for behavior and phenology. We outline challenges that currently limit the widespread application of biophysical models relating to data availability, training, and the lack of common software ecosystems. We also discuss progress and future developments that could allow these models to be applied to many species across large spatial extents and timeframes. Finally, we highlight how biophysical models are uniquely suited to solve global change biology problems that involve predicting and interpreting responses to environmental variability and extremes, multiple or shifting constraints, and novel abiotic or biotic environments., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Pollinating insects can be exposed to temperatures far from ambient air when visiting flowers, reducing their warming tolerance. Typically, such scenario occurs when flowers are exposed to solar radiation. The case of thermogenic flowers is particular because they warm up even when they are not exposed to solar energy. The flowers of Arum attract their pollinators with a deceptive method and trap them for a whole day, thereby imposing elevated temperature to visiting insects. Therefore, we predict a relatively high basal thermal tolerance in those insects. The aim of this study was to assess the thermal tolerance and warming tolerance of females of two fly species (genus Psychoda) pollinating Arum sp. (thermogenic plant). We measured their critical temperature (CTmax) and its response to rate of temperature increase as well as acclimation period to moderate temperature of 25 °C. We found relatively low CTmax (33.7 °C on average) for both species, and a weak response to acclimation period and ramping rate. In general, the thermal tolerance increased with a rapid ramping in temperature. To evaluate the warming tolerance, we compared thermal tolerance limits to flower temperatures measured in the field. We highlighted that the temperature of the thermogenic floral organ could reach values close to the thermal tolerance threshold of pollinators. This discovery raises questions about the sustainability of the interaction between these thermogenic plants and their pollinators., Competing Interests: Declaration of competing interest The authors declare no conflict of interest., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Historic approaches to understanding biological responses to climate change have viewed climate as something external that happens to organisms. Organisms, however, at least partially influence their own climate experience by moving within local mosaics of microclimates. Such behaviors are increasingly being incorporated into models of species distributions and climate sensitivity. Less attention has focused on how organisms alter microclimates via extended phenotypes: phenotypes that extend beyond the organismal surface, including structures that are induced or built. We argue that predicting the consequences of climate change for organismal performance and fitness will depend on understanding the expression and consequences of extended phenotypes, the microclimatic niches they generate, and the power of plasticity and evolution to shape those niches., Competing Interests: Declaration of interests No interests are declared., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Herbivore insects have strong impacts on leaf gas exchange when feeding on the plant. Leaf age also drives leaf gas exchanges but the interaction of leaf age and phloem herbivory has been largely underexplored. We investigated the amplitude and direction of herbivore impact on leaf gas exchange across a wide range of leaf age in the apple tree-apple green aphid ( Aphis pomi ) system. We measured the gas exchange (assimilation and transpiration rates, stomatal conductance and internal CO 2 concentration) of leaves infested versus non-infested by the aphid across leaf age. For very young leaves up to 15 days-old, the gas exchange rates of infested leaves were similar to those of non-infested leaves. After few days, photosynthesis, stomatal conductance and transpiration rate increased in infested leaves up to about the age of 30 days, and gradually decreased after that age. By contrast, gas exchanges in non-infested leaves gradually decreased across leaf age such that they were always lower than in infested leaves. Aphids were observed on relatively young leaves up to 25 days and despite the positive effect on leaf photosynthesis and leaf performance, their presence negatively affected the growth rate of apple seedlings. Indeed, aphids decreased leaf dry mass, leaf surface, and leaf carbon content except in old leaves. By contrast, aphids induced an increase in leaf nitrogen content and the deviation relative to non-infested leaves increased with leaf age. Overall, the impacts of aphids at multiple levels of plant performance depend on leaf age. While aphids cause an increase in some leaf traits (gas exchanges and nitrogen content), they also depress others (plant growth rate and carbon content). The balance between those effects, as modulated by leaf age, may be the key for herbivory mitigation in plants., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Pincebourde and Ngao.)

Temperature strongly drives physiological and ecological processes in ectotherms. While many species rely on behavioural thermoregulation to avoid thermal extremes, others build structures (nests) that confer a shelter against climate variability and extremes. However, the microclimate inside nests remains unknown for most insects. We investigated the thermal environment inside the nest of a temperate winter-developing insect species, the pine processionary moth (PPM), Thaumetopoea pityocampa. Gregarious larvae collectively build a silken nest at the beginning of the cold season. We tested the hypothesis that it provides a warmer microenvironment to larvae. First, we monitored temperature inside different types of nests varying in the number of larvae inside. Overall, nest temperature was positively correlated to global radiation and air temperature. At noon, when global radiation was maximal, nest temperature exceeded air temperature by up to 11.2-16.5 °C depending on nest type. In addition, thermal gradients of amplitude from 6.85 to 15.5 °C were observed within nests, the upper part being the warmest. Second, we developed a biophysical model to predict temperature inside PPM nests based on heat transfer equations and to explain this important temperature excess. A simple model version accurately predicted experimental measurements, confirming that nest temperature is driven mainly by radiation load. Finally, the model showed that nest temperature increases at the same rate as air temperature change. We conclude that some pest insects already live in warm microclimates by building their own sheltering nest. This effect should be considered when studying the impact of climate change on phenology and distribution., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Global change includes a substantial increase in the frequency and intensity of extreme high temperatures (EHTs), which influence insects at almost all levels. The number of studies showing the ecological importance of EHTs has risen in recent years, but the knowledge is rather dispersed in the contemporary literature. In this article, we review the biological and ecological effects of EHTs actually experienced in the field, i.e., when coupled to fluctuating thermal regimes. First, we characterize EHTs in the field. Then, we summarize the impacts of EHTs on insects at various levels and the processes allowing insects to buffer EHTs. Finally, we argue that the mechanisms leading to positive or negative impacts of EHTs on insects can only be resolved from integrative approaches considering natural thermal regimes. Thermal extremes, perhaps more than the gradual increase in mean temperature, drive insect responses to climate change, with crucial impacts on pest management and biodiversity conservation.

The SoilTemp database will identify the microhabitats that best buffer the amplitude of warming. The temperature heterogeneity at spatial scales below the meter also requires attention. A worldwide database of temperatures near any surface is still lacking. This article is a Commentary on Lembrechts et al., 26, 6616-6629., (© 2020 John Wiley & Sons Ltd.)

Climate warming impacts biological systems profoundly. Climatologists deliver predictions about warming amplitude at coarse scales. Nevertheless, insects are small, and it remains unclear how much of the warming at coarse scales appears in the microclimates where they live. We propose a simple method for determining the pertinent spatial scale of insect microclimates. Recent studies have quantified the ability of forest understory to buffer thermal extremes, but these microclimates typically are characterized at spatial scales much larger than those determined by our method. Indeed, recent evidence supports the idea that insects can be thermally adapted even to fine scale microclimatic patterns, which can be highly variable. Finally, we discuss how microhabitat surfaces may buffer or magnify the amplitude of climate warming., (Copyright © 2020. Published by Elsevier Inc.)

Vector-borne pathogens are known to alter the phenotypes of their primary hosts and vectors, with implications for disease transmission as well as ecology. Here we show that a plant virus, barley yellow dwarf virus, increases the surface temperature of infected host plants (by an average of 2 °C), while also significantly enhancing the thermal tolerance of its aphid vector Rhopalosiphum padi (by 8 °C). This enhanced thermal tolerance, which was associated with differential upregulation of three heat-shock protein genes, allowed aphids to occupy higher and warmer regions of infected host plants when displaced from cooler regions by competition with a larger aphid species, R. maidis. Infection thereby led to an expansion of the fundamental niche of the vector. These findings show that virus effects on the thermal biology of hosts and vectors can influence their interactions with one another and with other, non-vector organisms.

The thermal limit of ectotherms provides an estimate of vulnerability to climate change. It differs between contrasting microhabitats, consistent with thermal ecology predictions that a species' temperature sensitivity matches the microclimate it experiences. However, observed thermal limits may differ between ectotherms from the same environment, challenging this theory. We resolved this apparent paradox by showing that ectotherm activity generates microclimatic deviations large enough to account for differences in thermal limits between species from the same microhabitat. We studied upper lethal temperature, effect of feeding mode on plant gas exchange, and temperature of attacked leaves in a community of six arthropod species feeding on apple leaves. Thermal limits differed by up to 8 °C among the species. Species that caused an increase in leaf transpiration (+182%), thus cooling the leaf, had a lower thermal limit than those that decreased leaf transpiration (-75%), causing the leaf to warm up. Therefore, cryptic microclimatic variations at the scale of a single leaf determine the thermal limit in this community of herbivores. We investigated the consequences of these changes in plant transpiration induced by plant-insect feedbacks for species vulnerability to thermal extremes. Warming tolerance was similar between species, at ±2 °C, providing little margin for resisting increasingly frequent and intense heat waves. The thermal safety margin (the difference between thermal limit and temperature) was greatly overestimated when air temperature or intact leaf temperature was erroneously used. We conclude that feedback processes define the vulnerability of species in the phyllosphere, and beyond, to thermal extremes., Competing Interests: The authors declare no conflict of interest., (Copyright © 2019 the Author(s). Published by PNAS.)

Estimating leaf temperature distributions (LTDs) in canopies is crucial in forest ecology. Leaf temperature affects the exchange of heat, water, and gases, and it alters the performance of leaf-dwelling species such as arthropods, including pests and invaders. LTDs provide spatial variation that may allow arthropods to thermoregulate in the face of long-term changes in mean temperature or incidence of extreme temperatures. Yet, recording LTDs for entire canopies remains challenging. Here, we use an energy-exchange model (RATP) to examine the relative roles of climatic, structural, and physiological factors in influencing three-dimensional LTDs in tree canopies. A Morris sensitivity analysis of 13 parameters showed, not surprisingly, that climatic factors had the greatest overall effect on LTDs. In addition, however, structural parameters had greater effects on LTDs than did leaf physiological parameters. Our results suggest that it is possible to infer forest canopy LTDs from the LTDs measured or simulated just at the surface of the canopy cover over a reasonable range of parameter values. This conclusion suggests that remote sensing data can be used to estimate 3D patterns of temperature variation from 2D images of vegetation surface temperatures. Synthesis and applications . Estimating the effects of LTDs on natural plant-insect communities will require extending canopy models beyond their current focus on individual species or crops. These models, however, contain many parameters, and applying the models to new species or to mixed natural canopies depends on identifying the parameters that matter most. Our results suggest that canopy structural parameters are more important determinants of LTDs than are the physiological parameters that tend to receive the most empirical attention.

Understanding the effects of temperature on prey-predator interactions is a key issue to predict the response of natural communities to climate change. Higher temperatures are expected to induce an increase in predation rates. However, little is known on how temperature influences close-range encounter of prey-predator interactions, such as predator's attack velocities. Based on the speed-accuracy trade-off concept, we hypothesized that the increase in predator attack velocity by increasing temperature reduces the accuracy of the attack, leading to a lower probability of capture. We tested this hypothesis on the dragonfly larvae Anax imperator and the zooplankton prey Daphnia magna . The prey-predator encounters were video-recorded at high speed, and at three different temperatures. Overall, we found that (1) temperature had a strong effect on predator's attack velocities, (2) prey did not have the opportunity to move and/or escape due to the high velocity of the predator during the attack, and (3) neither velocity nor temperature had significant effects on the capture success. By contrast, the capture success mainly depended on the accuracy of the predator in capturing the prey. We found that (4) some 40% of mistakes were undershooting and some 60% aimed below or above the target. No lateral mistake was observed. These results did not support the speed-accuracy trade-off hypothesis. Further studies on dragonfly larvae with different morphological labial masks and speeds of attacks, as well as on prey with different escape strategies, would provide new insights into the response to environmental changes in prey-predator interactions.

Arthropods at the surface of plants live in particular microclimatic conditions that can differ from atmospheric conditions. The temperature of plant leaves can deviate from air temperature, and leaf temperature influences the eco-physiology of small insects. The activity of insects feeding on leaf tissues, may, however, induce changes in leaf surface temperatures, but this effect was only rarely demonstrated. Using thermography analysis of leaf surfaces under controlled environmental conditions, we quantified the impact of presence of apple green aphids on the temperature distribution of apple leaves during early infestation. Aphids induced a slight change in leaf surface temperature patterns after only three days of infestation, mostly due to the effect of aphids on the maximal temperature that can be found at the leaf surface. Aphids may induce stomatal closure, leading to a lower transpiration rate. This effect was local since aphids modified the configuration of the temperature distribution over leaf surfaces. Aphids were positioned at temperatures near the maximal leaf surface temperatures, thus potentially experiencing the thermal changes. The feedback effect of feeding activity by insects on their host plant can be important and should be quantified to better predict the response of phytophagous insects to environmental changes., Competing Interests: The authors declare no conflict of interest. The founding sponsors had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, and in the decision to publish the results.

Temperature is spatially heterogeneous over leaf surfaces, yet the underlying mechanisms are not fully resolved. We hypothesized that the 3D leaf microtopography determines locally the amount of incoming irradiation flux at leaf surface, thereby driving the temperature gradient over the leaf surface. This hypothesis was tested by developing a model of leaf temperature heterogeneity that includes the development of the leaf boundary layer, the microtopography of the leaf surface and the physiological response of the leaf. Temperature distributions under various irradiation loads (1) over apple leaves based on their 3D microtopography, (2) over simulated flat (2D) apple leaves and (3) over 3D leaves with a transpiration rate distributed as in 2D leaves were simulated. Accuracy of the predictions was quantified by comparing model outputs and thermographic measurements of leaf surface temperature under controlled conditions. Only the model with 3D leaves predicted accurately the spatial heterogeneity of surface temperature over single leaves, whereas the mean temperature was well predicted by both 2D and 3D leaves. We suggest that in these conditions, the 3D leaf microtopography is the primary driver of leaf surface heterogeneity in temperature when the leaf is exposed to a light/heat source., (© 2017 John Wiley & Sons Ltd.)

Most tropical ectotherms live near their physiological limits for temperature. Substantial ecological effects of global change are predicted in the tropics despite the low amplitude of temperature change. These predictions assume that tropical ectotherms experience air temperature as measured by weather stations or predicted by global circulation models. The body temperature of ectotherms, however, can deviate from ambient air when the organism samples the mosaic of microclimates at fine scales. The thermal heterogeneity of tropical landscapes has been quantified only rarely in comparison to temperate habitats, limiting our ability to infer the vulnerability to warming of tropical ectotherms. Here, we used thermal imaging to quantify the heterogeneity in surface temperatures across spatial scales, from the micro- up to landscape scale, at the top of an Inselberg in French Guiana. We measured the thermal heterogeneity at the scale of Clusia nemorosa leaves, by categorizing leaves in full sun versus leaves in the shade to quantify the microclimatic variance available to phytophagous insects. Then, we measured the thermal heterogeneity at the scales of the single shrub and the landscape, for several sites differing in their orientation toward the sun to quantify the microclimatic heterogeneity available for larger ectotherms. All measurements were made three times per day over four consecutive days. There was a high level of thermal heterogeneity at all spatial scales. The thermal variance varied between scales, increasing from the within-leaf surface to the landscape scale. It also shifted across the day in different ways depending on the spatial scale. Then, using a set of published data, we compared the critical temperature (CTmax) of neo-tropical ectotherms and temperature distributions. The portion of space above the CTmax varied substantially depending on spatial scale and taxa. Insects were particularly at risk at the surface of leaves exposed to solar radiation but not on shaded leaves. By contrast, ants tolerated elevated surface temperatures and can survive almost anywhere in the habitat. We suggest that the fine scale mosaic of microclimates in the tropics modulates the vulnerability of ectotherms to warming. By moving just a few meters, or even a few centimeters, small tropical ectotherms can radically change their microclimatic temperature and escape overheating., (© The Author 2016. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

When predicting the response of organisms to global change, models use measures of climate at a coarse resolution from general circulation models or from downscaled regional models. Organisms, however, do not experience climate at such large scales. The climate heterogeneity over a landscape and how much of that landscape an organism can sample will determine ultimately the microclimates experienced by organisms. This past few decades has seen an important increase in the number of studies reporting microclimatic patterns at small scales. This synthesis intends to unify studies reporting microclimatic heterogeneity (mostly temperature) at various spatial scales, to infer any emerging trends, and to discuss the causes and consequences of such heterogeneity for organismal performance and with respect to changing land use patterns and climate. First, we identify the environmental drivers of heterogeneity across the various spatial scales that are pertinent to ectotherms. The thermal heterogeneity at the local and micro-scales is mostly generated by the architecture or the geometrical features of the microhabitat. Then, the thermal heterogeneity experienced by individuals is modulated by behavior. Second, we survey the literature to quantify thermal heterogeneity from the micro-scale up to the scale of a landscape in natural habitats. Despite difficulties in compiling studies that differ much in their design and aims, we found that there is as much thermal heterogeneity across micro-, local and landscape scales, and that the temperature range is large in general (>9 °C on average, and up to 26 °C). Third, we examine the extent to which urban habitats can be used to infer the microclimatic patterns of the future. Urban areas generate globally drier and warmer microclimatic patterns and recent evidence suggest that thermal traits of ectotherms are adapted to them. Fourth, we explore the interplay between microclimate heterogeneity and the behavioral thermoregulatory abilities of ectotherms in setting their overall performance. We used a random walk framework to show that the thermal heterogeneity allows a more precise behavioral thermoregulation and a narrower temperature distribution of the ectotherm compared to less heterogeneous microhabitats. Finally, we discuss the potential impacts of global change on the fine scale mosaics of microclimates. The amplitude of change may differ between spatial scales. In heterogeneous microhabitats, the amplitude of change at micro-scale, caused by atmospheric warming, can be substantial while it can be limited at the local and landscape scales. We suggest that the warming signal will influence species performance and biotic interactions by modulating the mosaic of microclimates., (© The Author 2016. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Over the last few decades, biologists have made substantial progress in understanding relationships between changing climates and organism performance. Much of this work has focused on temperature because it is the best kept of climatic records, in many locations it is predicted to keep rising into the future, and it has profound effects on the physiology, performance, and ecology of organisms, especially ectothermic organisms which make up the vast majority of life on Earth. Nevertheless, much of the existing literature on temperature-organism interactions relies on mean temperatures. In reality, most organisms do not directly experience mean temperatures; rather, they experience variation in temperature over many time scales, from seconds to years. We propose to shift the focus more directly on patterns of temperature variation, rather than on means per se, and present a framework both for analyzing temporal patterns of temperature variation and for incorporating those patterns into predictions about organismal biology. In particular, we advocate using the Fourier transform to decompose temperature time series into their component sinusoids, thus allowing transformations between the time and frequency domains. This approach provides (1) standardized ways of visualizing the contributions that different frequencies make to total temporal variation; (2) the ability to assess how patterns of temperature variation have changed over the past half century and may change into the future; and (3) clear approaches to manipulating temporal time series to ask "what if" questions about the potential effects of future climates. We first summarize global patterns of change in temperature variation over the past 40 years; we find meaningful changes in variation at the half day to yearly times scales. We then demonstrate the utility of the Fourier framework by exploring how power added to different frequencies alters the overall incidence of long-term waves of high and low temperatures, and find that power added to the lowest frequencies greatly increases the probability of long-term heat and cold waves. Finally, we review what is known about the time scales over which organismal thermal performance curves change in response to variation in the thermal environment. We conclude that integrating information characterizing both the frequency spectra of temperature time series and the time scales of resulting physiological change offers a powerful new avenue for relating climate, and climate change, to the future performance of ectothermic organisms., (© The Author 2016. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Gas composition is an important component of any micro-environment. Insects, as the vast majority of living organisms, depend on O2 and CO2 concentrations in the air they breathe. Low O2 (hypoxia), and high CO2 (hypercarbia) levels can have a dramatic effect. For phytophagous insects that live within plant tissues (endophagous lifestyle), gas is exchanged between ambient air and the atmosphere within the insect habitat. The insect larva contributes to the modification of this environment by expiring CO2. Yet, knowledge on the gas exchange network in endophagous insects remains sparse. Our study identified mechanisms that modulate gas composition in the habitat of endophagous insects. Our aim was to show that the mere position of the insect larva within plant tissues could be used as a proxy for estimating risk of occurrence of hypoxia and hypercarbia, despite the widely diverse life history traits of these organisms. We developed a conceptual framework for a gas diffusion network determining gas composition in endophagous insect habitats. We applied this framework to mines, galls and insect tunnels (borers) by integrating the numerous obstacles along O2 and CO2 pathways. The nature and the direction of gas transfers depended on the physical structure of the insect habitat, the photosynthesis activity as well as stomatal behavior in plant tissues. We identified the insect larva position within the gas diffusion network as a predictor of risk exposure to hypoxia and hypercarbia. We ranked endophagous insect habitats in terms of risk of exposure to hypoxia and/or hypercarbia, from the more to the less risky as cambium mines>borer tunnels≫galls>bark mines>mines in aquatic plants>upper and lower surface mines. Furthermore, we showed that the photosynthetically active tissues likely assimilate larval CO2 produced. In addition, temperature of the microhabitat and atmospheric CO2 alter gas composition in the insect habitat. We predict that (i) hypoxia indirectly favors the evolution of cold-tolerant gallers, which do not perform well at high temperatures, and (ii) normoxia (ambient O2 level) in mines allows miners to develop at high temperatures. Little is known, however, about physiological and morphological adaptations to hypoxia and hypercarbia in endophagous insects. Endophagy strongly constrains the diffusion processes with cascading consequences on the evolutionary ecology of endophagous insects., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

We analyze the effects of changing patterns of thermal availability, in space and time, on the performance of small ectotherms. We approach this problem by breaking it into a series of smaller steps, focusing on: (1) how macroclimates interact with living and nonliving objects in the environment to produce a mosaic of thermal microclimates and (2) how mobile ectotherms filter those microclimates into realized body temperatures by moving around in them. Although the first step (generation of mosaics) is conceptually straightforward, there still exists no general framework for predicting spatial and temporal patterns of microclimatic variation. We organize potential variation along three axes-the nature of the objects producing the microclimates (abiotic versus biotic), how microclimates translate macroclimatic variation (amplify versus buffer), and the temporal and spatial scales over which microclimatic conditions vary (long versus short). From this organization, we propose several general rules about patterns of microclimatic diversity. To examine the second step (behavioral sampling of locally available microclimates), we construct a set of models that simulate ectotherms moving on a thermal landscape according to simple sets of diffusion-based rules. The models explore the effects of both changes in body size (which affect the time scale over which organisms integrate operative body temperatures) and increases in the mean and variance of temperature on the thermal landscape. Collectively, the models indicate that both simple behavioral rules and interactions between body size and spatial patterns of thermal variation can profoundly affect the distribution of realized body temperatures experienced by ectotherms. These analyses emphasize the rich set of problems still to solve before arriving at a general, predictive theory of the biological consequences of climate change., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Metabolic rate is a positive function of body weight, a rule valid for most organisms and the basis of several theories of metabolic ecology. For adult insects, however, the diversity of relationships between body mass and respiration remains unexplained. The aim of this study is to relate the respiratory metabolism of a parasitoid with body weight and foraging activity. We compared the metabolic rate of groups of starving and host-fed females of the parasitoid Eupelmus vuilleti recorded with respirometry for 7days, corresponding to the mean lifetime of starving females and over half of the lifetime of foraging females. The dynamics of carbohydrate, lipid and protein in the body of foraging females were quantified with biochemical techniques. Body mass and all body nutrients declined sharply from the first day onwards. By contrast, the CO2 produced and the O2 consumed increased steadily. Starving females showed the opposite trend, identifying foraging as the reason for the respiration increase of feeding females. Two complementary physiological processes explain the unexpected relationship between increasing metabolic rate and declining body weight. First, host hemolymph is a highly unbalanced food, and the excess nutrients (protein and carbohydrate) need to be voided, partially through excretion and partially through respiration. Second, a foraging young female produces eggs at an increasing rate during the first half of its lifetime, a process that also increases respiration. We posit that the time-varying metabolic rate contributions of the feeding and reproductive processes supplements the contribution of the structural mass and lead to the observed trend. We extend our explanations to other insect groups and discuss the potential for unification using Dynamic Energy Budget theory., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

The impact of warming on the persistence and distribution of ectotherms is often forecasted from their warming tolerance, inferred as the difference between their upper thermal limit and macroclimate temperature. Ectotherms, however, are thermally adapted to their microclimates, which can deviate substantially from macroscale conditions. Ignoring microclimates can therefore bias estimates of warming tolerance. We compared warming tolerance of an insect across its ontogeny when calculated from macro- and microclimate temperatures. We used a heat balance model to predict experienced microclimate temperatures from macroclimate, and we measured thermal limits for several life stages. The model shows a concomitant increase in microclimate temperatures and thermal limits across insect ontogeny, despite the fact that they all share the same macroclimate. Consequently, warming tolerance; as estimated from microclimate temperature, remained constant across ontogeny. When calculated from macroclimate temperature, however, warming tolerance was overestimated by 7-10 degrees C, depending on the life stage. Therefore, errors are expected when predicting persistence and distribution shifts of ectotherms in changing climates using macroclimate rather than microclimate.

Despite decades of work on climate change biology, the scientific community remains uncertain about where and when most species distributions will respond to altered climates. A major barrier is the spatial mismatch between the size of organisms and the scale at which climate data are collected and modeled. Using a meta-analysis of published literature, we show that grid lengths in species distribution models are, on average, ca. 10 000-fold larger than the animals they study, and ca. 1000-fold larger than the plants they study. And the gap is even worse than these ratios indicate, as most work has focused on organisms that are significantly biased toward large size. This mismatch is problematic because organisms do not experience climate on coarse scales. Rather, they live in microclimates, which can be highly heterogeneous and strongly divergent from surrounding macroclimates. Bridging the spatial gap should be a high priority for research and will require gathering climate data at finer scales, developing better methods for downscaling environmental data to microclimates, and improving our statistical understanding of variation at finer scales. Interdisciplinary collaborations (including ecologists, engineers, climatologists, meteorologists, statisticians, and geographers) will be key to bridging the gap, and ultimately to providing scientifically grounded data and recommendations to conservation biologists and policy makers., (© 2013 John Wiley & Sons Ltd.)

Body temperature is a more pertinent variable to physiological stress than ambient air temperature. Modeling and empirical studies on the impacts of climate change on ectotherms usually assume that body temperature within organisms is uniform. However, many ectotherms show significant within-body temperature heterogeneity. The relationship between regional heterothermy and the response of ectotherms to sublethal and lethal conditions remains underexplored. We quantified within-body thermal heterogeneity in an intertidal sea star (Pisaster ochraceus) during aerial exposure at low tide to examine the lethal and sublethal effects of temperatures of different body regions. In manipulative experiments, we measured the temperature of the arms and central disc, as well as survival and arm abscission under extreme aerial conditions. Survival was related strongly to central disc temperature. Arms were generally warmer than the central disc in individuals that survived aerial heating, but we found the reverse in those that died. When the central disc reached sublethal temperatures of 31-35°C, arms reached temperatures of 33-39°C, inducing arm abscission. The absolute temperature of individual arms was a poor predictor of arm abscission, but the arms lost were consistently the hottest at the within-individual scale. Therefore, the vital region of this sea star may remain below the lethal threshold under extreme conditions, possibly through water movement from the arms to the central disc and/or evaporative cooling, but at the cost of increased risk of arm abscission. Initiation of arm abscission seems to reflect a whole-organism response while death occurs as a result of stress acting directly on central disc tissues.

Climate warming experiments generally test the ecological effects of constant treatments while neglecting the influence of more realistic patterns of environmental fluctuations. Thus, little is known regarding how the temporal interaction between multiple episodes of thermal stress influences biotic interactions. We measured the sensitivity of predation rate in an intertidal sea star to changing levels of temporal coincidence of underwater and aerial thermal stress events. In laboratory trials, we controlled for intensity, variance and temporal patterning of both underwater and aerial body temperature. Predation rate decreased as underwater and aerial thermal stress episodes became temporally non-coincident, despite a similar intensity and variance among treatments. Experiments under constant conditions were a poor predictor of more complex environmental scenarios because of these strong temporal interactions. Such temporal interactions may be widespread in various ecosystems, suggesting a strong need for empirical studies and models that link environmental complexity, physiology, behaviour and species interactions., (© 2012 Blackwell Publishing Ltd/CNRS.)

The body temperature of ectotherms is influenced by the interaction of abiotic conditions, morphology, and behavior. Although organisms living in different thermal habitats may exhibit morphological plasticity or move from unfavorable locations, there are few examples of animals adjusting their thermal properties in response to short-term changes in local conditions. Here, we show that the intertidal sea star Pisaster ochraceus modulates its thermal inertia in response to prior thermal exposure. After exposure to high body temperature at low tide, sea stars increase the amount of colder-than-air fluid in their coelomic cavity when submerged during high tide, resulting in a lower body temperature during the subsequent low tide. Moreover, this buffering capacity is more effective when seawater is cold during the previous high tide. This ability to modify the volume of coelomic fluid provides sea stars with a novel thermoregulatory "backup" when faced with prolonged exposure to elevated aerial temperatures.

1. One major gap in our ability to predict the impacts of climate change is a quantitative analysis of temperatures experienced by organisms under natural conditions. We developed a framework to describe and quantify the impacts of local climate on the mosaic of microclimates and physiological states of insects within tree canopies. This approach was applied to a leaf mining moth feeding on apple leaf tissues. 2. Canopy geometry was explicitly considered by mapping the 3D position and orientation of more than 26 000 leaves in an apple tree. Four published models for canopy radiation interception, energy budget of leaves and mines, body temperature and developmental rate of the leaf miner were integrated. Model predictions were compared with actual microclimate temperatures. The biophysical model accurately predicted temperature within mines at different positions within the tree crown. 3. Field temperature measurements indicated that leaf and mine temperature patterns differ according to the regional climatic conditions (cloudy or sunny) and depending on their location within the canopy. Mines in the sun can be warmer than those in the shade by several degrees and the heterogeneity of mine temperature was incremented by 120%, compared with that of leaf temperature. 4. The integrated model was used to explore the impact of both warm and exceptionally hot climatic conditions recorded during a heat wave on the microclimate heterogeneity at canopy scale. During warm conditions, larvae in sunlight-exposed mines experienced nearly optimal growth conditions compared with those within shaded mines. The developmental rate was increased by almost 50% in the sunny microhabitat compared with the shaded location. Larvae, however, experienced optimal temperatures for their development inside shaded mines during extreme climatic conditions, whereas larvae in exposed mines were overheating, leading to major risks of mortality. 5. Tree canopies act as both magnifiers and reducers of the climatic regime experienced in open air outside canopies. Favourable and risky spots within the canopy do change as a function of the climatic conditions at the regional scale. The shifting nature of the mosaic of suitable and risky habitats may explain the observed uniform distribution of leaf miners within tree canopies.

Herbivory alters plant gas exchange but the effects depend on the type of leaf damage. In contrast to ectophagous insects, leaf miners, by living inside the leaf tissues, do not affect the integrity of the leaf surface. Thus, the effect of leaf miners on CO2 uptake and water-use efficiency by leaves remains unclear. We explored the impacts of the leaf-mining moth Phyllonorycter blancardella (Lepidoptera: Gracillariidae) on light responses of the apple leaf gas exchanges to determine the balance between the negative effects of reduced photosynthesis and potential positive impacts of increased water-use efficiency (WUE). Gas exchange in intact and mined leaf tissues was measured using an infrared gas analyser. The maximal assimilation rate was slightly reduced but the light response of net photosynthesis was not affected in mined leaf tissues. The transpiration rate was far more affected than the assimilation rate in the mine integument as a result of stomatal closure from moderate to high irradiance level. The WUE was about 200% higher in the mined leaf tissues than in intact leaf portions. Our results illustrate a novel mechanism by which plants might minimize losses from herbivore attacks; via trade-offs between the negative impacts on photosynthesis and the positive effects of increased WUE.

Very little is known about alterations in microclimate when an herbivore feeds on host plant. Modifications of leaf transmittance properties induced by feeding activity of the leaf miner Phyllonorycter blancardella F. were measured using a spectrometer. Their effects on the herbivore's body temperature and respiration rate have been determined under controlled conditions and varying radiation level employing an infrared gas analyser. By feeding within leaf tissues, a miner induces the formation of feeding windows which transmit a large portion of incoming radiations within a mine. As a result, body temperature and respiration rate increase with radiation level when positioned below feeding windows. Therefore, the miner is not always protected from radiations despite living within plant tissues. The amount of CO(2) released by larvae below feeding windows at high radiation levels is about five-fold that recorded in the dark. By contrast, body temperature and respiration rate increase only slightly with radiation level when the insect is positioned below intact tissues through which radiation is only weakly transmitted. A mine offers its inhabitant a heterogeneous light environment that allows the insect larva to thermoregulate through behavioural modification. Our results highlight the importance of physical feedbacks induced by herbivory which alter significantly an insect's metabolism independently of its nutritional state.