Tropical montane cloud forests (TMCFs) inhabit regions rich in biodiversity that play an important role in the local and regional water cycle. Canopy plants such as epiphytes and hemiepiphytes are an important component of the biodiversity in the TMCF and therefore play a significant role in the carbon, nutrient, and water cycles. With only partial or no access to resources on the ground, canopy plants may be vulnerable to changes in climate that increase canopy temperatures and decrease atmospheric humidity or precipitation inputs. Despite their importance in the TMCF, little is known about variation in functional strategies relating to drought avoidance or drought tolerance of canopy plants. In this study, we quantified variation in a number of functional traits in 11 species of epiphytes and hemiepiphytes in a Costa Rican TMCF. We also generated pressure–volume and xylem vulnerability curves that we used as indicators of drought tolerance. In addition, we hand‐sectioned fresh leaves and examined cross sections under a microscope to quantify leaf thickness, mesophyll thickness and the thickness of water storage cell layers (i.e., hydrenchyma), if present. Lastly, we determined the capacity for foliar water uptake in the laboratory and measured whole‐plant transpiration in the field. A trade‐off was found between traits that confer relative drought resistance and foliar water uptake capacity vs. traits that confer leaf capacitance and relative drought avoidance. This trade‐off may represent an additional axis of the leaf economics spectrum that is unique to epiphytes. We also found that all species had the capacity for foliar uptake of water and that this process contributed substantially to their water balance. On average, foliar uptake of water contributed to the reabsorption of 70% of the water transpired over a relatively wet, 34‐day study period. Our results indicate that canopy plants can mitigate water loss substantially via internal water storage or that they can directly utilize cloud water to offset losses. Our results indicate that species that rely on foliar uptake of water may be more vulnerable to projected changes in climate than species that buffer the effects of drought via internal water storage.

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.

Epiphyte communities comprise important components of many forest ecosystems in terms of biomass and diversity, but little is known regarding trade-offs that underlie diversity and structure in these communities or the impact that microclimate has on epiphyte trait allocation. We measured 22 functional traits in vascular epiphyte communities across six sites that span a microclimatic gradient in a tropical montane cloud forest region in Costa Rica. We quanti ed traits that relate to carbon and nitrogen allocation, gas exchange, water storage, and drought tolerance. Functional diversity was high in all but the lowest elevation site where drought likely limits the success of certain species with particular trait combinations. For most traits, variation was explained by relationships with other traits (trait covariance), rather than differences in microclimate across sites. Although there were signi cant differences in microclimate, epiphyte abundance, and diversity, we found substantial overlap in multivariate trait space across ve of the sites. We found signi cant correlations between functional traits, many of which related to water storage, drought tolerance, and carbon allocation. This suite of trait correlations suggests that the epiphyte community has evolved functional strategies along a drought avoidance versus drought tolerance continuum where leaf succulence emerged as a pivotal overall trait.

Epiphyte communities comprise important components of many forest ecosystems in terms of biomass and diversity, but little is known regarding trade-offs that underlie diversity and structure in these communities or the impact that microclimate has on epiphyte trait allocation. We measured 22 functional traits in vascular epiphyte communities across six sites that span a microclimatic gradient in a tropical montane cloud forest region in Costa Rica. We quantified traits that relate to carbon and nitrogen allocation, gas exchange, water storage, and drought tolerance. Functional diversity was high in all but the lowest elevation site where drought likely limits the success of certain species with particular trait combinations. For most traits, variation was explained by relationships with other traits (trait co-variance), rather than differences in microclimate across sites. Although there were significant differences in microclimate, epiphyte abundance, and diversity, we found substantial overlap in multivariate trait space across five of the sites. We found significant correlations between functional traits, many of which related to water storage, drought tolerance, and carbon allocation. This suite of trait correlations suggests that the epiphyte community has evolved functional strategies along a drought avoidance versus drought tolerance continuum where leaf succulence emerged as a pivotal overall trait.