Shower thoughts: why scientists should spend more time in the rain.

Stormwater is a vital resource and dynamic driver of terrestrial ecosystem processes. However, processes controlling interactions during and shortly after storms are often poorly seen and poorly sensed when direct observations are substituted with technological ones. We discuss how human observations complement technological ones and the benefits of scientists spending more time in the storm. Human observation can reveal ephemeral storm-related phenomena such as biogeochemical hot moments, organismal responses, and sedimentary processes that can then be explored in greater resolution using sensors and virtual experiments. Storm-related phenomena trigger lasting, oversized impacts on hydrologic and biogeochemical processes, organismal traits or functions, and ecosystem services at all scales. We provide examples of phenomena in forests, across disciplines and scales, that have been overlooked in past research to inspire mindful, holistic observation of ecosystems during storms. We conclude that technological observations alone are insufficient to trace the process complexity and unpredictability of fleeting biogeochemical or ecological events without the shower thoughts produced by scientists' human sensory and cognitive systems during storms.

Drought decreases water storage capacity of two arboreal epiphytes with differing ecohydrological traits.

Arboreal epiphytes, plants that grow on trees, can significantly increase rainwater storage and evaporation (i.e., "interception") within canopies. Drought conditions may affect this hydrological role, as epiphytes' physiological responses change leaf properties that affect water retention. Drought-induced changes in epiphyte water storage capacity could substantially alter canopy hydrology, but have not been studied. We tested the effects of drought on the water storage capacity (Smax) of leaves and leaf properties of two epiphytes with distinct ecohydrological traits: resurrection fern (Pleopeltis polypodioides), and Spanish moss (Tillandsia usneoides). Both species are common in maritime forests of the Southeastern USA, where climate change is expected to decrease precipitation in spring and summer. To simulate drought, we dried leaves to 75 %, 50 %, and ~25 % of fresh weight, and quantified their Smax in fog chambers. We measured relevant leaf properties: hydrophobicity, minimum leaf conductance (gmin; a measure of water loss under drought), and Normalized Difference Vegetative Index (NDVI). We found that drought significantly reduced Smax and increased leaf hydrophobicity for both species, indicating that lower Smax may be due to shedding of droplets. While the overall reduction in Smax did not differ between the two species, they exhibited distinct drought responses. Dehydrated T. usneoides leaves had lower gmin, demonstrating the ability to limit water loss under drought. P. polypodioides increased gmin when dehydrated, consistent with its extraordinary ability to withstand water loss. NDVI decreased with dehydration in T. usneoides but not P. polypodioides. Our results suggest that increased drought may have a dramatic effect on canopy water cycling by reducing the Smax of epiphytes. Reduced rainfall interception and storage in forest canopies could have widespread effects on hydrological cycling, thus understanding the potential feedbacks of plant drought response on hydrology is crucial. This study highlights the importance of connecting foliar-scale plant response with broader hydrological processes.

Variation in cloud immersion, not precipitation, drives leaf trait plasticity and water relations in vascular epiphytes during an extreme drought.

PREMISE: Epiphytes are abundant in ecosystems such as tropical montane cloud forests where low-lying clouds are often in contact with vegetation. Climate projections for these regions include more variability in rainfall and an increase in cloud base heights, which would lead to drier conditions in the soil and atmosphere. While recent studies have examined the effects of drought on epiphytic water relations, the influence that atmospheric moisture has, either alone or in combination with drought, on the health and performance of epiphyte communities remains unclear. METHODS: We conducted a 10-week drought experiment on seven vascular epiphyte species in two shadehouses, one with warmer and drier conditions and another that was cooler and more humid. We measured water relations across control and drought-treatment groups and assessed functional traits of leaves produced during drought conditions to evaluate trait plasticity. RESULTS: Epiphytes exposed to drought and drier atmospheric conditions had a significant reduction in stomatal conductance and leaf water potential and an increase in leaf dry matter. Nonsucculent epiphytes from the drier shadehouse had the greatest shifts in functional traits, whereas succulent epiphytes released stored leaf water to maintain water status. CONCLUSIONS: Individuals in the drier shadehouse had a substantial reduction in performance, whereas drought-treated individuals that experienced cloud immersion displayed minimal changes in water status. Our results indicate that projected increases in the cloud base height will reduce growth and performance of epiphytic communities and that nonsucculent epiphytes may be particularly vulnerable.

Correction to: Allocation to leaf area and sapwood area affects water relations of co-occurring savanna and forest trees.

The original version of this article unfortunately contained a mistake. The Electronic supplementary material (ESM) was accompanying this article by mistake.

Foliar water uptake: Processes, pathways, and integration into plant water budgets.

Nearly all plant families, represented across most major biomes, absorb water directly through their leaves. This phenomenon is commonly referred to as foliar water uptake. Recent studies have suggested that foliar water uptake provides a significant water subsidy that can influence both plant water and carbon balance across multiple spatial and temporal scales. Despite this, our mechanistic understanding of when, where, how, and to what end water is absorbed through leaf surfaces remains limited. We first review the evidence for the biophysical conditions necessary for foliar water uptake to occur, focusing on the plant and atmospheric water potentials necessary to create a gradient for water flow. We then consider the different pathways for uptake, as well as the potential fates of the water once inside the leaf. Given that one fate of water from foliar uptake is to increase leaf water potentials and contribute to the demands of transpiration, we also provide a quantitative synthesis of observed rates of change in leaf water potential and total fluxes of water into the leaf. Finally, we identify critical research themes that should be addressed to effectively incorporate foliar water uptake into traditional frameworks of plant water movement.

Variation in the resilience of cloud forest vascular epiphytes to severe drought.

Epiphytes are common in tropical montane cloud forests (TMCFs) and play many important ecological roles, but the degree to which these unique plants will be affected by changes in climate is unknown. We investigated the drought responses of three vascular epiphyte communities bracketing the cloud base during a severe, El Niño-impacted dry season. Epiphytes were instrumented with sap flow probes in each site. Leaf water potential and pressure-volume curve parameters were also measured before and during the drought. We monitored the canopy microclimate in each site to determine the drivers of sap velocity across the sites. All plants greatly reduced their water use during the drought, but recovery occurred more quickly for plants in the lower and drier sites. Plants in drier sites also exhibited the greatest shifts in the osmotic potential at full saturation and the turgor loss point. Although all individuals survived this intense drought, epiphytes in the cloud forest experienced the slowest recovery, suggesting that plants in the TMCF are particularly sensitive to severe drought. Although vapor pressure deficit was an important driver of sap velocity in the highest elevation site, other factors, such as the volumetric water content of the canopy soil, were more important at lower (and warmer) sites.

Vapor pressure deficit predicts epiphyte abundance across an elevational gradient in a tropical montane region.

PREMISE OF THE STUDY: Tropical Montane Cloud Forests (TMCFs) are important ecosystems to study and preserve because of their high biodiversity and critical roles in local and regional ecosystem processes. TMCFs may be particularly affected by changes in climate because of the narrow bands of microclimate they occupy and the vulnerability of TMCF species to projected increases in cloud base heights and drought. A comprehensive understanding of the structure and function of TMCFs is lacking and difficult to attain because of variation in topography within and across TMCF sites. This causes large differences in microclimate and forest structure at both large and small scales. METHODS: In this study, we estimated the abundance of the entire epiphyte community in the canopy (bryophytes, herbaceous vascular plants, woody epiphytes, and canopy dead organic matter) in six sites. In each of the sites we installed a complete canopy weather station to link epiphyte abundance to a number of microclimatic parameters. KEY RESULTS: We found significant differences in epiphyte abundance across the sites; epiphyte abundance increased with elevation and leaf wetness, but decreased as vapor pressure deficit (VPD) increased. Epiphyte abundance had the strongest relationship with VPD; there were differences in VPD that could not be explained by elevation alone. CONCLUSIONS: By measuring this proxy of canopy VPD, TMCF researchers will better understand differences in microclimate and plant community composition across TMCF sites. Incorporating such information in comparative studies will allow for more meaningful comparisons across TMCFs and will further conservation and management efforts in this ecosystem.

Habitat moisture is an important driver of patterns of sap flow and water balance in tropical montane cloud forest epiphytes.

Microclimate in the tropical montane cloud forest (TMCF) is variable on both spatial and temporal scales and can lead to large fluctuations in both leaf-level transpiration and whole plant water use. While variation in transpiration has been found in TMCFs, the influence of different microclimatic drivers on plant water relations in this ecosystem has been relatively understudied. Within the TMCF, epiphytes may be particularly affected by natural variation in microclimate due to their partial or complete disassociation from soil resources. In this study, we examined the effects of seasonal microclimate on whole plant water balance in epiphytes in both an observational and a manipulative experiment. We also evaluated the effects of different microclimatic drivers using three hierarchical linear (mixed) models. On average, 31 % of total positive sap flow was recovered via foliar water uptake (FWU) over the course of the study. We found that precipitation was the greatest driver of foliar water uptake and nighttime sap flow in our study species and that both VPD and precipitation were important drivers to daytime sap flow. We also found that despite adaptations to withstand seasonal drought, an extended dry period caused severe desiccation in most plants despite a large reduction in leaf-level and whole plant transpiration. Our results indicate that the epiphytes studied rely on FWU to maintain positive water balance in the dry season and that increases in dry periods in the TMCF may be detrimental to these common members of the epiphyte community.

Water relations and microclimate around the upper limit of a cloud forest in Maui, Hawai'i.

The goal of this study was to determine the effects of atmospheric demand on both plant water relations and daily whole-tree water balance across the upper limit of a cloud forest at the mean base height of the trade wind inversion in the tropical trade wind belt. We measured the microclimate and water relations (sap flow, water potential, stomatal conductance, pressure-volume relations) of Metrosideros polymorpha Gaudich. var. polymorpha in three habitats bracketing the cloud forest's upper limit in Hawai'i to understand the role of water relations in determining ecotone position. The subalpine shrubland site, located 100 m above the cloud forest boundary, had the highest vapor pressure deficit, the least amount of rainfall and the highest levels of nighttime transpiration (EN) of all three sites. In the shrubland site, on average, 29% of daily whole-tree transpiration occurred at night, while on the driest day of the study 50% of total daily transpiration occurred at night. While EN occurred in the cloud forest habitat, the proportion of total daily transpiration that occurred at night was much lower (4%). The average leaf water potential (Ψleaf) was above the water potential at the turgor loss point (ΨTLP) on both sides of the ecotone due to strong stomatal regulation. While stomatal closure maintained a high Ψleaf, the minimum leaf water potential (Ψleafmin) was close to ΨTLP, indicating that drier conditions may cause drought stress in these habitats and may be an important driver of current landscape patterns in stand density.

Foggy days and dry nights determine crown-level water balance in a seasonal tropical Montane cloud forest.

The ecophysiology of tropical montane cloud forest (TMCF) trees is influenced by crown-level microclimate factors including regular mist/fog water inputs, and large variations in evaporative demand, which in turn can significantly impact water balance. We investigated the effect of such microclimatic factors on canopy ecophysiology and branch-level water balance in the dry season of a seasonal TMCF in Veracruz, Mexico, by quantifying both water inputs (via foliar uptake, FU) and outputs (day- and night-time transpiration, NT). Measurements of sap flow, stomatal conductance, leaf water potential and pressure-volume relations were obtained in Quercus lanceifolia, a canopy-dominant tree species. Our results indicate that FU occurred 34% of the time and led to the recovery of 9% (24 ± 9.1 L) of all the dry-season water transpired from individual branches. Capacity for FU was independently verified for seven additional common tree species. NT accounted for approximately 17% (46 L) of dry-season water loss. There was a strong correlation between FU and the duration of leaf wetness events (fog and/or rain), as well as between NT and the night-time vapour pressure deficit. Our results show the clear importance of fog and NT for the canopy water relations of Q. lanceifolia.

Ecological thresholds at the savanna-forest boundary: how plant traits, resources and fire govern the distribution of tropical biomes.

Fire shapes the distribution of savanna and forest through complex interactions involving climate, resources and species traits. Based on data from central Brazil, we propose that these interactions are governed by two critical thresholds. The fire-resistance threshold is reached when individual trees have accumulated sufficient bark to avoid stem death, whereas the fire-suppression threshold is reached when an ecosystem has sufficient canopy cover to suppress fire by excluding grasses. Surpassing either threshold is dependent upon long fire-free intervals, which are rare in mesic savanna. On high-resource sites, the thresholds are reached quickly, increasing the probability that savanna switches to forest, whereas low-resource sites are likely to remain as savanna even if fire is infrequent. Species traits influence both thresholds; saplings of savanna trees accumulate bark thickness more quickly than forest trees, and are more likely to become fire resistant during fire-free intervals. Forest trees accumulate leaf area more rapidly than savanna trees, thereby accelerating the transition to forest. Thus, multiple factors interact with fire to determine the distribution of savanna and forest by influencing the time needed to reach these thresholds. Future work should decipher multiple environmental controls over the rates of tree growth and canopy closure in savanna.

Allocation to leaf area and sapwood area affects water relations of co-occurring savanna and forest trees.

Water availability is a principal factor limiting the distribution of closed-canopy forest in the seasonal tropics, suggesting that forest tree species may not be well adapted to cope with seasonal drought. We studied 11 congeneric species pairs, each containing one forest and one savanna species, to test the hypothesis that forest trees have a lower capacity to maintain seasonal homeostasis in water relations relative to savanna species. To quantify this, we measured sap flow, leaf water potential (Psi(L)), stomatal conductance (g (s)), wood density, and Huber value (sapwood area:leaf area) of the 22 study species. We found significant differences in the water relations of these two species types. Leaf area specific hydraulic conductance of the soil/root/leaf pathway (G (t)) was greater for savanna species than forest species. The lower G (t) of forest trees resulted in significantly lower Psi(L) and g (s) in the late dry season relative to savanna trees. The differences in G (t) can be explained by differences in biomass allocation of savanna and forest trees. Savanna species had higher Huber values relative to forest species, conferring greater transport capacity on a leaf area basis. Forest trees have a lower capacity to maintain homeostasis in Psi(L) due to greater allocation to leaf area relative to savanna species. Despite significant differences in water relations, relationships between traits such as wood density and minimum Psi(L) were indistinguishable for the two species groups, indicating that forest and savanna share a common axis of water-use strategies involving multiple traits.

Tree topkill, not mortality, governs the dynamics of savanna-forest boundaries under frequent fire in central Brazil.

Tropical savanna and forest are recognized to represent alternate stable states, primarily determined by feedbacks with fire. Vegetation-fire dynamics in each of these vegetation types are largely determined by the influence of the vegetation on fire behavior, as well as the effects of fire behavior on tree mortality, topkill (defined here as complete death of the aerial biomass, regardless of whether the plant recovers by resprouting), and rate of growth of resprouts. We studied the effect of fire on three savanna-forest boundaries in central Brazil. Fire intensity was greater in savanna than forest, as inferred by a twofold greater height of stem charring. Despite lower fire intensity, forest tree species exhibited higher rates of topkill, which was best explained by their thinner bark, relative to savanna species. Following topkill, there was no tendency for sprouts of savanna trees to grow faster than those of forest species, contrary to expectations, nor was whole-plant mortality higher in forest than in savanna. This contrasts with observations of high rates of postburn mortality in many other tropical forests. The low tree mortality in these transitional forests suggests that the dynamic of these natural savanna-forest boundaries is fundamentally different from that of forest boundaries originating from deforestation in the humid tropics. The forests studied here appear to be much more resilient to occasional incursion of fire from the savanna, despite being unable to invade frequently burned savanna. The thin bark of forest species makes them particularly susceptible to the "fire trap," whereby repeated topkill of small trees prevents recruitment into adult size classes. Rapid growth will be particularly important for forest species to escape the fire trap, so we predict that, where fire is frequent, forests should be restricted to high-resource sites. Here, Mg2+ and Ca2+ concentrations had particularly strong effects on postburn growth rates, suggesting that these elements may most strongly limit the distribution of forest in these fire-prone savannas.