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.

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Forest microclimates and climate change: Importance, drivers and future research agenda.

Average body temperatures (Tb) of summer-active Sceloporus merriami were lower in the morning (33.30C) than in the evening (37.0?) at an arid locality in Big Bend National Park, Texas. During midafternoon, lizards were inactive and retreated to deep crevices and caves. According to a multivariate analysis, overall rates of lizard activities (movements, feeding, and social displays) were similar between morning and evening activity, indicating that lizards were behaving similarly during the different activity periods. However, at 370, estimated rates of metabolism and water loss were greater and sprint speed was lower relative to these rates at 330, suggesting that substantial resource and performance costs may have been imposed by the observed higher evening activity Tb. To investigate this unusual pattern of body temperature variation, I compared the morning and evening distributions of available operative environmental temperatures (Te) as estimated by an array of randomly deployed and oriented copper models of S. merriami placed on rock and ground surfaces within lizard home ranges. In the morning, lizards had access to a wide range of Te (300-600), and a large fraction of the Te distribution was below 360 at the time of peak morning activity. During the morning, a hypothetical increase in Tb to within the interval 36?-38? would have allowed lizards to occupy 900% depending on the exact time), and microclimates below 360 were unavailable until just before dark. The available Te distributions in the evening forced lizards to accept a higher activity Tb, and concomitantly higher metabolic and water loss rates, and lower locomotor performance, or remain inactive in crevices and caves. Thus in the evening, and not in the morning, the gain in usable home range due to accepting a higher Tb was great, and the lizards exploited this fraction at an increased Tb only in the evening. These data suggest that the distribution of available Te can directly influence the cost/benefit economics of lizard body temperature selection.

Trade-offs in activity time and physiological performance for thermoregulating desert

Gloger's rule is an ecogeographical rule that links animal colouration with climatic variation. This rule is named after C.W.L. Gloger who was one of the first to summarise the associations between climatic variation and animal colouration, noting in particular that birds and mammals seemed more pigmented in tropical regions. The term 'Gloger's rule' was coined by B. Rensch in 1929 and included different patterns of variation from those described by Gloger. Rensch defined the rule in two ways: a simple version stating that endothermic animals are predicted to be darker in warmer and humid areas due to the increased deposition of melanin pigments; and a complex version that includes the differential effects of humidity and temperature on both main types of melanin pigments - eu- and phaeo-melanin. The blackish eu-melanins are predicted to increase with humidity, and decrease only at extreme low temperatures, while the brown-yellowish phaeomelanins prevail in dry and warm regions and decrease rapidly with lower temperatures. A survey of the literature indicates that there is considerable variation/confusion in the way Gloger's rule is understood (based on 271 studies that define the rule). Whereas the complex version is hardly mentioned, only a quarter of the definitions are consistent with the simple version of Gloger's rule (darker where warm and wet), and most definitions mention only the effects of humidity (darker where wet). A smaller subset of studies define the rule based on other correlated climatic and environmental variables such as vegetation, latitude, altitude, solar radiation, etc., and a few even contradict the original definition (darker where cold). Based on the literature survey, I synthesised the qualitative (N = 124 studies) and quantitative (meta-analytically, N = 38 studies, 241 effects) evidence testing the simple version of Gloger's rule (I found no tests of the complex version). Both lines of evidence supported the predicted effects of humidity (and closely linked variables) on colour variation, but not the effects of temperature. Moreover, humidity effects are not restricted to birds and mammals, as the data indicate that these effects also apply to insects. This suggests that the simple version of Gloger's rule as originally defined may not be valid, and possibly that the rule should be re-formulated in terms of humidity effects only. I suggest, however, that more data are needed before such a reformulation, due to potential publication biases. In conclusion, I recommend that authors cite Rensch when referring to Gloger's rule and that they make clear which version they are referring to. Future research should concentrate on rigorously testing the validity and generality of both versions of Gloger's rule and establishing the mechanism(s) responsible for the patterns it describes. Since humidity seems to be the core climatic variable behind Gloger's rule, I suggest that the two most plausible mechanisms are camouflage and protection against parasites/pathogens, the latter possibly through pleiotropic effects on the immune system. Understanding the processes that lead to climatic effects on animal colouration may provide insights into past and future patterns of adaptation to climatic change.

A review of Gloger's rule, an ecogeographical rule of colour: definitions, interpretations and evidence.

Gloger's rule is usually interpreted as predicting darker coloured animals in warmer and more humid/vegetated regions. The relative importance of temperature and rainfall or vegetation is however unclear, and often only one variable is tested at a time, mainly through proxies. Here, I assess the predictions of Gloger's rule for interspecific achromatic plumage variation (dark to light variation) for an entire avifauna (551 species of Australian landbirds). I tested the effects of climatic variables (temperature and rainfall) and vegetation structure on plumage reflectance at species and assemblage level (100x100 km cells), controlling for phylogenetic relatedness and spatial autocorrelation. To assess the robustness of these results I compared observed results with those of a null distribution of effects obtained from repeatedly simulating random plumage reflectance evolution on the phylogeny. At both the species and assemblage level, darker coloured birds were found in wetter and colder regions and in more densely vegetated habitats. Simulations confirm results at the species level and the effect of temperature at the assemblage level, but rainfall and vegetation effects at the assemblage level fall within the distribution of simulated effects and should be interpreted with care. Interspecific colour variation in Australian birds supports Gloger's rule for rainfall/vegetation, but shows the opposite pattern for temperature. Darker colours in wet and vegetated environments are consistent with the role of melanin pigmentation in preventing feather degradation by bacteria, but also with background-matching for camouflage. Darker plumage might be beneficial in colder regions or detrimental in warmer regions if it affects thermoregulation, a selective force often only assumed to be of importance for ectotherms. The data highlight the need to test the generality of biogeographic rules across levels and at broad scale. Experimental work is needed to confirm the mechanisms linking plumage achromatic variation to climate.

This article is protected by copyright. All rights reserved.

https://www.onlinelibrary.wiley.com/doi/pdf/10.1111/ecog.03040

Darker where cold and wet: Australian birds follow their own version of Gloger's rule

Ecogeographical rules that associate climate with organismal form and function can reveal patterns of climatic adaptation. Two rules link animal coloration with climate: Gloger's rule (darker coloration where wet and warm), and Bogert's rule (darker coloration where cold). Whereas Gloger's rule was proposed for endotherms, and Bogert's rule for ectotherms, both rules may apply more broadly, despite their seemingly opposing effects. Here, we test this contradiction on a global scale across passerine birds. Consistent with Gloger's rule, birds were darker in wetter areas and, following Bogert's rule, lighter where warm, although birds became lighter again at very low temperatures. Rainfall and temperature had antagonistic or additive effects depending on their pattern of covariation, and this predicted whether birds followed the rules. We integrate both rules into a general framework to explain heterogeneity in climatic effects on coloration, which has implications to understand patterns of diversification, climatic adaptation and climate change impacts.

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Reconciling ecogeographical rules: rainfall and temperature predict global colour variation in the largest bird radiation

Morphology mediates the relationship between an organism's body temperature and its environment. Dark organisms, for example, tend to absorb heat more quickly than lighter individuals, which could influence their responses to temperature. Therefore, temperature-related traits such as morphology may affect patterns of species abundance, richness, and community assembly across a broad range of spatial scales. In this study, we examined variation in color lightness and body size within butterfly communities across hot and cool habitats in the tropical woodland–rainforest ecosystems of northeast Queensland, Australia. Using thermal imaging, we documented the absorption of solar radiation relative to color lightness and wingspan and then built a phylogenetic tree based on available sequences to analyze the effects of habitat on these traits within a phylogenetic framework. In general, darker and larger individuals were more prevalent in cool, closed-canopy rainforests than in immediately adjacent and hotter open woodlands. In addition, darker and larger butterflies preferred to be active in the shade and during crepuscular hours, while lighter and smaller butterflies were more active in the sun and midday hours—a pattern that held after correcting for phylogeny. Our ex situ experiment supported field observations that dark and large butterflies heated up faster than light and small butterflies under standardized environmental conditions. Our results show a thermal consequence of butterfly morphology across habitats and how environmental factors at a microhabitat scale may affect the distribution of species based on these traits. Furthermore, this study highlights how butterfly species might differentially respond to warming based on ecophysiological traits and how thermal refuges might emerge at microclimatic and habitat scales.

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Chin Cheung Tang

Papers

Cool habitats support darker and bigger butterflies in Australian tropical forests.