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Plot-based monitoring has yielded much information on the taxonomic diversity and carbon (C) storage in tropical lowland forests of the Amazon basin. This has resulted in an improved understanding of the relationship between lowland forest biomass dynamics and global change drivers, such as climate change and atmospheric CO2 concentration. Much less attention has been paid to the mountain ecosystems of South America that comprise montane forests and alpine vegetation (páramo, puna, high Andean grasslands, wetlands, and alpine heath). This vegetation complex provides a variety of ecosystem services and forms a natural laboratory along various physiographic, geological and evolutionary history/biogeography, and land use history gradients. Here, we review existing empirical understanding and model-based approaches to quantify the contribution of mountain ecosystems to ecosystem service provision in the rapidly changing socioecological setting of the South American mountains. The objective of this paper is to outline a broad road map for the implementation of mountain vegetation into dynamic global vegetation models (DGVM) for use in Earth System Models (ESM), based on our current understanding of their structure and function and of their responsiveness to global change drivers. We also identify treeline processes, critical in mountain ecosystems, as key missing elements in DGVMs/ESMs, and thus explore in addition a treeline model. Stocktaking of the availability of empirical data was undertaken from eight research sites along the Andes and in south-eastern Brazil. Out of eight sites, two (one each in Venezuela and Brazil) had some climate, ecological and ecophysiological data potentially suitable to parametrise a DGVM. Tree biomass data were available for six sites. A preliminary assessment of the Joint UK Land Environment Simulator (JULES) DGVM was made to identify gaps in available data and their impacts on model parametrisation and calibration. Additionally, the potential climate-determined elevation of the treeline was modelled to check the DGVM for its ability to identify the transition between the montane forest and alpine vegetation. Outcomes of the evaluation of the JULES land surface model identified the following key processes in montane forests: temperature-related decrease in net primary production, respiration, and allocation to above-ground biomass and increase in soil C stocks with elevation. There was a variable agreement between simulated biomass and those derived from field measurements via allometric equations. We identified major gaps between data availability and the needs for process-based modelling of South American mountain vegetation and its dynamics in DGVMs. To bridge this gap, we propose a transdisciplinary network, composed of members of the theoretical/modelling and empirical scientific communities, to study the natural dynamics of mountain ecosystems and their responses to global change drivers locally, regionally and at the continental scale, within a social-ecological system framework. The work presented here forms the basis for the design of data collection from field measurements and instrumental monitoring stations to parametrise and verify DGVMs. The network is designed to collaborate with and complement existing long-term research initiatives in the region and will adopt existing standard field protocols. Complementary protocols will ensure compatibility between field data collection and data needed for process-based and empirical models. [ABSTRACT FROM AUTHOR]

Summary: Tropical ecosystems have the highest levels of biodiversity, cycle more water and absorb more carbon than any other terrestrial ecosystem on Earth. Consequently, these ecosystems are extremely important components of Earth's climatic system and biogeochemical cycles. Plant hydraulics is an essential discipline to understand and predict the dynamics of tropical vegetation in scenarios of changing water availability. Using published plant hydraulic data we show that the trade‐off between drought avoidance (expressed as deep‐rooting, deciduousness and capacitance) and hydraulic safety (P50 – the water potential when plants lose 50% of their maximum hydraulic conductivity) is a major axis of physiological variation across tropical ecosystems. We also propose a novel and independent axis of hydraulic trait variation linking vulnerability to hydraulic failure (expressed as the hydraulic safety margin (HSM)) and growth, where inherent fast‐growing plants have lower HSM compared to slow‐growing plants. We surmise that soil nutrients are fundamental drivers of tropical community assembly determining the distribution and abundance of the slow‐safe/fast‐risky strategies. We conclude showing that including either the growth‐HSM or the resistance‐avoidance trade‐off in models can make simulated tropical rainforest communities substantially more vulnerable to drought than similar communities without the trade‐off. These results suggest that vegetation models need to represent hydraulic trade‐off axes to accurately project the functioning and distribution of tropical ecosystems. [ABSTRACT FROM AUTHOR]

Ecologists seek a general scheme to classify the diversity of plant responses to environmental factors into a few strategies (e.g. competitor—C, stress tolerant—S, ruderal—R), while plant physiologists seek a mechanistic scheme to explain such different responses (e.g. tolerance, escape, avoidance). So far, few attempts have been made to combine both perspectives into plant eco‐physiological strategies. Moreover, the relative contribution of different strategies to maintain both community structure and ecosystem functioning during drought has rarely been assessed. Thus, limiting our capacity to predict how extreme events caused by climate change will affect plant communities.Here, we present an integrated framework to identify plant eco‐physiological strategies and to estimate their contribution to community originality (diversity of trait combinations), dominance (species relative frequency) and ecosystem functioning (productivity and evapotranspiration).We applied this framework in a tropical montane grassland in Brazil (Campos de Altitude) and found three eco‐physiological strategies co‐occurring in this community (S‐tolerance/avoidance, CS‐escape/tolerance and CR‐escape/avoidance). While CS‐species contributed more to dominance and functionality, CR‐ and S‐species contributed more to originality. Therefore, all three strategies were important to support the grassland form and function.Synthesis. Plants exhibit different strategies, as well as different contributions to community and ecosystem attributes. We developed an integrated approach to both identify strategies and estimate their relative contribution. Thereby, as droughts intensify, we can better predict which plants are more likely to be lost and how their loss will impact the communities and ecosystem where they occur. This knowledge is necessary for specifying conservation priorities and for developing more efficient conservation practices. [ABSTRACT FROM AUTHOR]

Summary: Land surface models (LSMs) typically use empirical functions to represent vegetation responses to soil drought. These functions largely neglect recent advances in plant ecophysiology that link xylem hydraulic functioning with stomatal responses to climate.We developed an analytical stomatal optimization model based on xylem hydraulics (SOX) to predict plant responses to drought. Coupling SOX to the Joint UK Land Environment Simulator (JULES) LSM, we conducted a global evaluation of SOX against leaf‐ and ecosystem‐level observations.SOX simulates leaf stomatal conductance responses to climate for woody plants more accurately and parsimoniously than the existing JULES stomatal conductance model. An ecosystem‐level evaluation at 70 eddy flux sites shows that SOX decreases the sensitivity of gross primary productivity (GPP) to soil moisture, which improves the model agreement with observations and increases the predicted annual GPP by 30% in relation to JULES. SOX decreases JULES root‐mean‐square error in GPP by up to 45% in evergreen tropical forests, and can simulate realistic patterns of canopy water potential and soil water dynamics at the studied sites.SOX provides a parsimonious way to incorporate recent advances in plant hydraulics and optimality theory into LSMs, and an alternative to empirical stress factors. See also the Commentary on this article by Anderegg & Venturas, 226: 1535–1538. [ABSTRACT FROM AUTHOR]

The current generation of dynamic global vegetation models (DGVMs) lacks a mechanistic representation of vegetation responses to soil drought, impairing their ability to accurately predict Earth system responses to future climate scenarios and climatic anomalies, such as El Niño events. We propose a simple numerical approach to model plant responses to drought coupling stomatal optimality theory and plant hydraulics that can be used in dynamic global vegetation models (DGVMs). The model is validated against stand-scale forest transpiration (E) observations from a long-term soil drought experiment and used to predict the response of three Amazonian forest sites to climatic anomalies during the twentieth century. We show that our stomatal optimization model produces realistic stomatal responses to environmental conditions and can accurately simulate how tropical forest E responds to seasonal, and even long-term soil drought. Our model predicts a stronger cumulative effect of climatic anomalies in Amazon forest sites exposed to soil drought during El Niño years than can be captured byalternativeempiricaldrought representation schemes. The contrasting responses between our model and empirical drought factors highlight theutilityofhydraulically-basedstomatal optimizationmodels to represent vegetation responses to drought and climatic anomalies inDGVMs. This article is part of a discussion meeting issue 'The impact of the 2015/ 2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'. [ABSTRACT FROM AUTHOR]

Aim: The Anthropocene climate crisis may shift the distribution range of various species. Global Climate Models predict an increase in temperature and changes in precipitation across the Brazilian tropical semiarid region. Based on the joint analysis of functional traits and plant distribution models, we aim to identify which functional traits define the vulnerability of plants to climate change. Location: Caatinga, Brazil. Taxon: Trees‐shrub plants. Methods: We selected 36 species common to at least 25% of the floristic and phytosociological surveys made in the Brazilian semiarid region. We modelled both the current and future spatial distribution of these species and investigated whether functional traits, such as plant height, leaf area, leaf thickness, specific leaf area, stem specific density, and seed size can explain the range shifts of these species. Results: Models indicate that 58.4% of the species are expected to expand their distribution range in the future. These species tend to be shorter and produce large seeds and thin leaves, which confer higher hydraulic safety and resistance of seeds to prolonged droughts. Meanwhile, 41.6% of the species are expected to reduce their distribution ranges in the future. Most of these are characterized as tall individuals with thick leaves and small seeds. Conclusion: Species with traits that confer resistance to drought are likely to increase distribution towards the coast and to areas where the Caatinga borders with other domains. Also, these species with low height, thin leaves and large seeds may expand their distribution to the slopes and plateaus. The retraction of taller plants and with thicker leaves may have profound implications for the structure and functioning of Brazilian semiarid ecosystems, given their role in ecosystem processes and carbon cycle. Retractions can result in a reduction in the number of local species, which will impact regional biodiversity. [ABSTRACT FROM AUTHOR]

Many tropical montane cloud forest (TMCF) trees are capable of foliar water uptake (FWU) during leaf-wetting events. In this study, we tested the hypothesis that maintenance of leaf turgor during periods of fog exposure and soil drought is related to species' FWU capacity., We conducted several experiments using apoplastic tracers, deuterium labeling and leaf immersion in water to evaluate differences in FWU among three common TMCF tree species. We also measured the effect of regular fog exposure on the leaf water potential of plants subjected to soil drought and used these data to model species' response to long-term drought., All species were able to absorb water through their leaf cuticles and/or trichomes, although the capacity to do so differed between species. During the drought experiment, the species with higher FWU capacity maintained leaf turgor for a longer period when exposed to fog, whereas the species with lower FWU exerted tighter stomatal regulation to maintain leaf turgor. Model results suggest that without fog, species with high FWU are more likely to lose turgor during seasonal droughts., We show that leaf-wetting events are essential for trees with high FWU, which tend to be more anisohydric, maintaining leaf turgor during seasonal droughts. [ABSTRACT FROM AUTHOR]

Trees from tropical montane cloud forest (TMCF) display very dynamic patterns of water use. They are capable of downwards water transport towards the soil during leaf-wetting events, likely a consequence of foliar water uptake (FWU), as well as high rates of night-time transpiration (Enight) during drier nights. These two processes might represent important sources of water losses and gains to the plant, but little is known about the environmental factors controlling these water fluxes. We evaluated how contrasting atmospheric and soil water conditions control diurnal, nocturnal and seasonal dynamics of sap flow in Drimys brasiliensis (Miers), a common Neotropical cloud forest species. We monitored the seasonal variation of soil water content, micrometeorological conditions and sap flow of D. brasiliensis trees in the field during wet and dry seasons. We also conducted a greenhouse experiment exposing D. brasiliensis saplings under contrasting soil water conditions to deuterium-labelled fog water. We found that during the night D. brasiliensis possesses heightened stomatal sensitivity to soil drought and vapour pressure deficit, which reduces night-time water loss. Leaf-wetting events had a strong suppressive effect on tree transpiration (E). Foliar water uptake increased in magnitude with drier soil and during longer leaf-wetting events. The difference between diurnal and nocturnal stomatal behaviour in D. brasiliensis could be attributed to an optimization of carbon gain when leaves are dry, as well as minimization of nocturnal water loss. The leaf-wetting events on the other hand seem important to D. brasiliensis water balance, especially during soil droughts, both by suppressing tree transpiration (E) and as a small additional water supply through FWU. Our results suggest that decreases in leaf-wetting events in TMCF might increase D. brasiliensis water loss and decrease its water gains, which could compromise its ecophysiological performance and survival during dry periods. [ABSTRACT FROM AUTHOR]

Background Tropical montane cloud forests (TMCFs) are characterized by a unique set of biological and hydroclimatic features, including frequent and/or persistent fog, cool temperatures, and high biodiversity and endemism. These forests are one of the most vulnerable ecosystems to climate change given their small geographic range, high endemism and dependence on a rare microclimatic envelope. The frequency of atmospheric water deficits for some TMCFs is likely to increase in the future, but the consequences for the integrity and distribution of these ecosystems are uncertain. In order to investigate plant and ecosystem responses to climate change, we need to know how TMCF species function in response to current climate, which factors shape function and ecology most and how these will change into the future. Scope This review focuses on recent advances in ecophysiological research of TMCF plants to establish a link between TMCF hydrometeorological conditions and vegetation distribution, functioning and survival. The hydraulic characteristics of TMCF trees are discussed, together with the prevalence and ecological consequences of foliar uptake of fog water (FWU) in TMCFs, a key process that allows efficient acquisition of water during cloud immersion periods, minimizing water deficits and favouring survival of species prone to drought-induced hydraulic failure. Conclusions Fog occurrence is the single most important microclimatic feature affecting the distribution and function of TMCF plants. Plants in TMCFs are very vulnerable to drought (possessing a small hydraulic safety margin), and the presence of fog and FWU minimizes the occurrence of tree water deficits and thus favours the survival of TMCF trees where such deficits may occur. Characterizing the interplay between microclimatic dynamics and plant water relations is key to foster more realistic projections about climate change effects on TMCF functioning and distribution. [ABSTRACT FROM PUBLISHER]

Foliar water uptake ( FWU) is a common water acquisition mechanism for plants inhabiting temperate fog-affected ecosystems, but the prevalence and consequences of this process for the water and carbon balance of tropical cloud forest species are unknown., We performed a series of experiments under field and glasshouse conditions using a combination of methods (sap flow, fluorescent apoplastic tracers and stable isotopes) to trace fog water movement from foliage to belowground components of Drimys brasiliensis. In addition, we measured leaf water potential, leaf gas exchange, leaf water repellency and growth of plants under contrasting soil water availabilities and fog exposure in glasshouse experiments to evaluate FWU effects on the water and carbon balance of D. brasiliensis saplings., Fog water diffused directly through leaf cuticles and contributed up to 42% of total foliar water content. FWU caused reversals in sap flow in stems and roots of up to 26% of daily maximum transpiration. Fog water transported through the xylem reached belowground pools and enhanced leaf water potential, photosynthesis, stomatal conductance and growth relative to plants sheltered from fog., Foliar uptake of fog water is an important water acquisition mechanism that can mitigate the deleterious effects of soil water deficits for D. brasiliensis. [ABSTRACT FROM AUTHOR]

Occult precipitation events (fog, dew and light rain) can alter plant water and nutritional status, both directly through the aerial uptake of surface water and nutrients, and indirectly via redistribution of atmospheric resources to the soil. However, current frameworks that explain niche segregation, species interactions and coexistence still consider that ground‐rooted plants obtain resources almost exclusively via root absorption from soil.Here, we expand the plant hydrological niches model to incorporate both soil and atmospheric resource‐axes, thus providing a more complete picture of how ground‐rooted terrestrial plants obtain, remobilise, share and compete for water and soluble nutrients.First, we describe how plants with different water acquisition strategies access directly or indirectly atmospheric resources. Then, we discuss how the use of such resources may promote spatiotemporal niche segregation, contributing to shape species distribution and abundance within plant communities. We illustrate this argument with examples from arid, mesic and wet vegetation types. Finally, we examine how climate and land‐use changes may influence plant hydrological niches, potentially altering community structure.Synthesis. Understanding how available atmospheric resources influences niche segregation in plant communities is a crucial step towards better predictions of species responses (e.g. changes in distribution, abundance and interactions) to climate change. [ABSTRACT FROM AUTHOR]

Tropical montane cloud forests (TMCF) have unique climatic conditions, which allow the coexistence of plant lineages with different phytogeographical origins from tropical versus temperate climates. Future climate projections suggest TMCFs will be subjected to increasing drought stress due to fog uplift and higher temperatures, possibly leading to tree mortality and local extinctions, and consequently changes in forest composition and functioning. Characterizing community functional composition, trade‐offs among traits and the drivers of community assembly is of utmost importance to improve our capacity to predict the response of montane plant communities to forecast climate change.Here, we aimed to test whether species from different phytogeographical origins (i.e. tropical – evergreen × deciduous − and temperate) differ in drought vulnerability and how the coexistence of these groups change the hydraulic composition of TMCFs. We used a framework based on measurements of key hydraulic traits (i.e. xylem embolism resistance, hydraulic safety margin, stomata control, turgor loss point, minimum water potential) of 16 dominant species (>70% of the forest basal area) within a TMCF in the Atlantic Rain Forest Domain in southeast Brazil. We used community‐weighted means to model whether removing each species group would change the community hydraulic functional composition.Temperate, tropical deciduous and tropical evergreen groups differ in their hydraulic functioning and these differences explain forest functional composition and taxa dominance. Temperate and tropical deciduous taxa were consistently more vulnerable hydraulically (i.e. lower safety margins and embolism resistance). The coexistence of different phytogeographical lineages is a key determinant of TMCF hydraulic composition. We also used models including phylogeny to evaluate the variation of hydraulic traits across phytogeographical groups, and the results suggest some niche conservatism associated with plant hydraulic functioning.Our results provide evidence of the importance of species phytogeographical origin on TMCF functioning, and niche conservatism in the evolution of hydraulic traits. The higher drought vulnerability observed in temperate group might be a mechanistic explanation for the restriction of temperate taxa distribution to wetter places during past colder and drier climate. Thus, we suggest hydraulic functional traits may be useful to predict future dynamics of TMCFs under changing climatic conditions. [ABSTRACT FROM AUTHOR]

Highlights • We quantify the fog regime of a Tropical Cloud Forest in high temporal resolution. • Fog occurrence is highly variable at all temporal scales and prevalent at night. • Fog strongly contributes to ecosystem water inputs. • Cloud regime, and not fog regime, dominates light and microclimatic variability. • Understanding cloud forest functioning may require high resolution studies of fog regimes. Abstract Fog is a frequent phenomenon in tropical montane cloud forests (TMCFs). These ecosystems are important to the water supply of adjacent lowland regions which is largely determined by the effect of fog on TCMF evapotranspiration rates and hydrological balance. Understanding fog regimes at fine-grained temporal resolution is key to predict plant functioning and effects of climatic changes in TMCFs, especially on key hydrological services that these forests provide. Here, we combine a suite of micrometeorological and hydrological sensors with a visibilimeter, a reliable sensor of fog occurrence, to gather fine-grained information on fog frequency, duration and timing and its contribution to water inputs, light availability and microclimatic variability in a Brazilian TMCF. Despite occurring on 64% of days, fog was highly variable at daily and seasonal scales, occurring mostly at night and during the rainy season. Approximately 1200 liters of fog were intercepted per tree per year (259 mm or 10.7% of total net precipitation). Fog also increased net precipitation provided by concomitant fog-rain events. Monthly net precipitation to precipitation ratio, a measure of how much water arrives at the soil and how much evaporates or is intercepted by the canopy, was 0.96 - much higher than the 0.72 typical of lowland rainforest, due to the additional fog water input on TMCF. Cloudiness, and not fog, dominated light availability and inter-day microclimatic variability (air temperature and vapor pressure deficit). High fog regime variability indicates that understanding TMCFs functioning requires integration of plant function with fine-grained data of fog and cloud occurrence. We discuss possible consequences of our results to TMCFs plant functioning. [ABSTRACT FROM AUTHOR]

In Brazil, the campos rupestres occur over the Brazilian shield, and are characterized by acidic nutrient-impoverished soils, which are particularly low in phosphorus (P). Despite recognition of the campos rupestres as a global biodiversity hotspot, little is known about the diversity of P-acquisition strategies and other aspects of plant mineral nutrition in this region., To explore nutrient-acquisition strategies and assess aspects of plant P nutrition, we measured leaf P and nitrogen (N) concentrations, characterized root morphology and determined the percentage arbuscular mycorrhizal ( AM) colonization of 50 dominant species in six communities, representing a gradient of soil P availability. Leaf manganese (Mn) concentration was measured as a proxy for carboxylate-releasing strategies., Communities on the most P-impoverished soils had the highest proportion of nonmycorrhizal ( NM) species, the lowest percentage of mycorrhizal colonization, and the greatest diversity of root specializations. The large spectrum of leaf P concentration and variation in root morphologies show high functional diversity for nutritional strategies. Higher leaf Mn concentrations were observed in NM compared with AM species, indicating that carboxylate-releasing P-mobilizing strategies are likely to be present in NM species., The soils of the campos rupestres are similar to the most P-impoverished soils in the world. The prevalence of NM strategies indicates a strong global functional convergence in plant mineral nutrition strategies among severely P-impoverished ecosystems. [ABSTRACT FROM AUTHOR]