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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.

Climate change is altering the dynamics, structure and function of the Amazon, a biome deeply connected to the Earth’s carbon cycle. Climate factors that control the spatial and temporal variations in forest photosynthesis have been well studied, but the influence of forest height and age on this controlling effect has rarely been considered. Here, we present remote sensing observations of solar-induced fluorescence (a proxy for photosynthesis), precipitation, vapour-pressure deficit and canopy height, together with estimates of forest age and aboveground biomass. We show that photosynthesis in tall Amazonian forests, that is, forests above 30 m, is three times less sensitive to precipitation variability than in shorter (less than 20 m) forests. Taller Amazonian forests are also found to be older, have more biomass and deeper rooting systems1, which enable them to access deeper soil moisture and make them more resilient to drought. We suggest that forest height and age are an important control of photosynthesis in response to interannual precipitation fluctuations. Although older and taller trees show less sensitivity to precipitation variations, they are more susceptible to fluctuations in vapour-pressure deficit. Our findings illuminate the response of Amazonian forests to water stress, droughts and climate change.

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.

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.

The functional trait approach has, as a central tenet, that plant traits are functional and shape individual performance, but this has rarely been tested in the field. Here, we tested the individual-based trait approach in a hyperdiverse Amazonian tropical rainforest and evaluated intraspecific variation in trait values, plant strategies at the individual level, and whether traits are functional and predict individual performance. We evaluated > 1300 tree saplings belonging to > 383 species, measured 25 traits related to growth and defense, and evaluated the effects of environmental conditions, plant size, and traits on stem growth. A total of 44% of the trait variation was observed within species, indicating a strong potential for acclimation. Individuals showed two strategy spectra, related to tissue toughness and organ size vs leaf display. In this nutrient- and light-limited forest, traits measured at the individual level were surprisingly poor predictors of individual growth performance because of convergence of traits and growth rates. Functional trait approaches based on individuals or species are conceptually fundamentally different: the species-based approach focuses on the potential and the individual-based approach on the realized traits and growth rates. Counterintuitively, the individual approach leads to a poor prediction of individual performance, although it provides a more realistic view on community dynamics.

1. The relationship between rooting depth and above-ground hydraulic traits can potentially define drought resistance strategies that are important in determining species distribution and coexistence in seasonal tropical forests, and understanding this is important for predicting the effects of future climate change in these ecosystems. 2. We assessed the rooting depth of 12 dominant tree species (representing c. 42% of the forest basal area) in a seasonal Amazon forest using the stable isotope ratios (δ¹⁸ and δ²H) of water collected from tree xylem and soils from a range of depths. We took advantage of a major ENSO-related drought in 2015/2016 that caused substantial evaporative isotope enrichment in the soil and revealed water mum dry season leaf water potential both in a normal year (2014; Ψnon-ENSO) and in an extreme drought year (2015; Ψnon-ENSO Furthermore, we measured xylem hydraulic traits that indicate water potential thresholds trees tolerate without risking hydraulic failure (P₅₀ and P₈₈). 3. We demonstrate that coexisting trees are largely segregated along a single hydrological niche axis defined by root depth differences, access to light and tolerance of low water potential. These differences in rooting depth were strongly related to tree size; diameter at breast height (DBH) explained 72% of the variation in the tree size; diameter at breast height (DBH) explained 72% of the variation in the δ¹⁸Oxylem Additionally, δ¹⁸Oxylem explained 49% of the variation in P₅₀ and 70% of P₈₈, with shallow-rooted species more tolerant of low water potentials, while δ¹⁸ of xylem water explained 47% and 77% of the variation of minimum Ψnon-ENSO and ΨENSO. 4. We propose a new formulation to estimate an effective functional rooting depth, i.e. the likely soil depth from which roots can sustain water uptake for physiological functions, using DBH as predictor of root depth at this site. Based on these estimates, we conclude that rooting depth varies systematically across the most abundant families, genera and species at the Tapajós forest, and that understorey species in particular are limited to shallow rooting depths. 5. Our results support the theory of hydrological niche segregation and its underlying trade-off related to drought resistance, which also affect the dominance structure of trees in this seasonal eastern Amazon forest. 6. Synthesis. Our results support the theory of hydrological niche segregation an demonstrate its underlying trade-off related to drought resistance (access to deep water vs. tolerance of very low water potentials). We found that the single hydrological axis defining water use traits was strongly related to tree size, and infer that periodic extreme droughts influence community composition and the dominance structure of trees in this seasonal eastern Amazon forest.

• The mangrove Avicennia marina adjusts internal salt concentrations by foliar salt secretion. Deliquescence of accumulated salt causes leaf wetting that may provide a water source for salt-secreting plants in arid coastal wetlands where high nocturnal humidity can usually support deliquescence whereas rainfall events are rare. We tested the hypotheses that salt deliquescence on leaf surfaces can drive top-down rehydration, and that such absorption of moisture from unsaturated atmospheres makes a functional contribution to dry season shoot water balances. • Sap flow and water relations were monitored to assess the uptake of atmospheric water by branches during shoot wetting events under natural and manipulated microclimatic conditions. • Reverse sap flow rates increased with increasing relative humidity from 70% to 89%, consistent with function of salt deliquescence in harvesting moisture from unsaturated atmospheres. Top-down rehydration elevated branch water potentials above those possible from root water uptake, subsidising transpiration rates and reducing branch vulnerability to hydraulic failure in the subsequent photoperiod. • Absorption of atmospheric moisture harvested through deliquescence of salt on leaf surfaces enhances water balances of Avicennia marina growing in hypersaline wetlands under arid climatic conditions. Top-down rehydration from these frequent, low intensity wetting events contributes to prevention of carbon starvation and hydraulic failure during drought.

Predictions of the magnitude and timing of leaf phenology in Amazonian forests remain highly controversial. Here, we use terrestrial LiDAR surveys every two weeks spanning wet and dry seasons in Central Amazonia to show that plant phenology varies strongly across vertical strata in old-growth forests, but is sensitive to disturbances arising from forest fragmentation. In combination with continuous microclimate measurements, we find that when maximum daily temperatures reached 35 °C in the latter part of the dry season, the upper canopy of large trees in undisturbed forests lost plant material. In contrast, the understory greened up with increased light availability driven by the upper canopy loss, alongside increases in solar radiation, even during periods of drier soil and atmospheric conditions. However, persistently high temperatures in forest edges exacerbated the upper canopy losses of large trees throughout the dry season, whereas the understory in these light-rich environments was less dependent on the altered upper canopy structure. Our findings reveal a strong influence of edge effects on phenological controls in wet forests of Central Amazonia. Even evergreen tropical forests can have seasonal dynamics, which may be sensitive to disturbance. Here, the authors combine high-resolution remote sensing observations and microclimate data to show that forest fragmentation impacts canopy phenology dynamics in the Amazon forest.

Plants allocate nutrients to specific leaf cell types, with commelinoid monocots preferentially allocating phosphorus (P) to the mesophyll and calcium (Ca) to the epidermis, whereas the opposite is thought to occur in eudicots. However, Proteaceae from severely P-impoverished habitats present the same P-allocation pattern as monocots. This raises the question of whether preferential P allocation to mesophyll cells is a phylogenetically conserved trait, exclusive to commelinoid monocots and a few Proteaceae, or a trait that has evolved multiple times to allow plants to cope with very low soil P availability. We analysed the P-allocation patterns of 16 species from 10 genera, eight families and six orders within three major clades of eudicots across different P-impoverished environments in Australia and Brazil, using elemental X-ray mapping to quantitatively determine leaf cell-specific nutrient concentrations. Many of the analysed species showed P-allocation patterns that differed substantially from that expected for eudicots. Instead, P-allocation patterns were strongly associated with the P availability in the natural habitat of the species, suggesting a convergent evolution of P-allocation patterns at the cellular level, with P limitation as selective pressure and without a consistent P-allocation pattern within eudicots. Here, we show that most eudicots from severely P-impoverished environments preferentially allocated P to their mesophyll. We surmise that this preferential P allocation to photosynthetically active cells might contribute to the very high photosynthetic P-use efficiency of species adapted to P-impoverished habitats.

• Species distribution is strongly driven by local and global gradients in water availability but the underlying mechanisms are not clear. Vulnerability to xylem embolism (P50) is a key trait that indicates how species cope with drought and might explain plant distribution patterns across environmental gradients. Here we address its role on species sorting along a hydrotopographical gradient in a central Amazonian rainforest and examine its variance at the community scale. • We measured P50 for 28 tree species, soil properties and estimated the hydrological niche of each species using an indicator of distance to the water table (HAND). • We found a large hydraulic diversity, covering as much as 44% of the global angiosperm variation in P50. We show that P50: contributes to species segregation across a hydrotopographic gradient in the Amazon, and thus to species coexistence; is the result of repeated evolutionary adaptation within closely related taxa; is associated with species tolerance to P-poor soils, suggesting the evolution of a stress-tolerance syndrome to nutrients and drought; and is higher for trees in the valleys than uplands. • The large observed hydraulic diversity and its association with topography has important implications for modelling and predicting forest and species resilience to climate change.