Z. Carter Berry

Cloud immersion: an important water source for spruce and fir saplings in the southern Appalachian Mountains

Cloud immersion can provide a potentially important moisture subsidy to plants in areas of frequent fog including the threatened spruce-fir communities of the southern Appalachian Mountains (USA). These mountaintop communities grow only above ~1,500xa0m elevation, harbor the endemic Abies fraseri, and have been proposed to exist because of frequent cloud immersion. While several studies have demonstrated the importance of cloud immersion to plant water balance, no study has evaluated the proportion of plant water derived from cloud moisture in this ecosystem. Using the isotopic mixing model, IsoSource, we analyzed the isotopic composition of hydrogen and oxygen for water extracted from ground water, deep soil, shallow soil, fog, and plant xylem at the upper and lower elevational limits both in May (beginning of the growing season) and October (end of the growing season). Cloud-immersion water contributed up to 31xa0% of plant water at the upper elevation sites in May. High-elevation plants of both species also experienced greater cloud immersion and had greater cloud water absorption (14–31xa0%) compared to low-elevation plants (4–17xa0%). Greater cloud water uptake occurred in May compared to October, despite similar rainfall and cloud-immersion frequencies. These results demonstrate the important water subsidy that cloud-immersion water can provide. With a warming climate leading potentially to increases in the ceiling of the cloud base and, thus, less frequent cloud immersion, persistence of these relic mountaintop forests may depend on the magnitude of these changes and the compensating capabilities of other water sources.

Ecophysiological importance of cloud immersion in a relic spruce–fir forest at elevational limits, southern Appalachian Mountains, USA

Climate warming predicts changes to the frequency and height of cloud-immersion events in mountain communities. Threatened southern Appalachian spruce–fir forests have been suggested to persist because of frequent periods of cloud immersion. These relic forests exist on only seven mountaintop areas, grow only above ca. 1,500xa0m elevation (maximum 2,037xa0m), and harbor the endemic Abies fraseri. To predict future distribution, we examined the ecophysiological effects of cloud immersion on saplings of A. fraseri and Picea rubens at their upper and lower elevational limits. Leaf photosynthesis, conductance, transpiration, xylem water potentials, and general abiotic variables were measured simultaneously on individuals at the top (1,960xa0m) and bottom (1,510xa0m) of their elevation limits on numerous clear and cloud-immersed days throughout the growing season. The high elevation sites had 1.5 as many cloud-immersed days (75xa0% of days) as the low elevation sites (56xa0% of days). Cloud immersion resulted in higher photosynthesis, leaf conductance, and xylem water potentials, particularly during afternoon measurements. Leaf conductance remained higher throughout the day with corresponding increases in photosynthesis and transpiration, despite low photon flux density levels, leading to an increase in water potentials from morning to afternoon. The endemic A. fraseri had a greater response in carbon gain and water balance in response to cloud immersion. Climate models predict warmer temperatures with a decrease in the frequency of cloud immersion for this region, leading to an environment on these peaks similar to elevations where spruce–fir communities currently do not exist. Because spruce–fir communities may rely on cloud immersion for improved carbon gain and water conservation, an upslope shift is likely if cloud ceilings rise. Their ultimate survival will likely depend on the magnitude of changes in cloud regimes.

Experimental cloud immersion and foliar water uptake in saplings of Abies fraseri and Picea rubens

Key messageFrequent cloud immersion events result in direct uptake of cloud water and improve plant water potentials during daylight hours in saplings of two dominant cloud forest species.AbstractIn ecosystems with frequent cloud immersion, the influence on plant water balance can be important. While cloud immersion can reduce plant water loss via transpiration, recent advances in methodology have suggested that many species also absorb water directly into leaves (foliar water uptake). The current study examines foliar water uptake and its influence on daily plant water balance in tree species of the endangered spruce–fir forest of the southern Appalachian Mountains, USA. These mountain-top communities are considered relic, boreal forests that may have persisted because of the benefits of frequent cloud immersion. We examined changes in needle water content, xylem water potentials, and stable isotope values in saplings of the two dominant tree species, Abies fraseri and Picea rubens before and after a 24xa0h period of experimental cloud immersion. Both species exhibited foliar water uptake following immersion, evidenced by substantial changes in stable isotope values of extracted needle water that reflected the composition of the fog water. In addition, total needle water content improved 3.7–6.4xa0% following experimental submersion and xylem water potentials were significantly greater (up to 0.33xa0MPa) in cloud-immersed plants over control plants. These results indicate that foliar water uptake may be an adaptive strategy for utilizing cloud water and improving overall tree vigor in these most southerly distributed boreal species.

Foliar uptake, carbon fluxes and water status are affected by the timing of daily fog in saplings from a threatened cloud forest.

In cloud forests, foliar uptake (FU) of water has been reported for numerous species, possibly acting to relieve daily water and carbon stress. While the prevalence of FU seems common, how daily variation in fog timing may affect this process has not been studied. We examined the quantity of FU, water potentials, gas exchange and abiotic variation at the beginning and end of a 9-day exposure to fog in a glasshouse setting. Saplings of Abies fraseri (Pursh) Poir. and Picea rubens Sarg. were exposed to morning (MF), afternoon (AF) or evening fog (EF) regimes to assess the ability to utilize fog water at different times of day and after sustained exposure to simulated fog. The greatest amount of FU occurred during MF (up to 50%), followed by AF (up to 23%) and then EF, which surprisingly had no FU. There was also a positive relationship between leaf conductance and FU, suggesting a role of stomata in FU. Moreover, MF and AF lead to the greatest improvements in daily water balance and carbon gain, respectively. Foliar uptake was important for improving plant ecophysiology but was influenced by diurnal variation in fog. With climate change scenarios predicting changes to cloud patterns and frequency that will likely alter diurnal patterns, cloud forests that rely on this water subsidy could be affected.

Co-occurring woody species have diverse hydraulic strategies and mortality rates during an extreme drought: Belowground hydraulic failure during drought

From 2011 to 2013, Texas experienced its worst drought in recorded history. This event provided a unique natural experiment to assess species-specific responses to extreme drought and mortality of four co-occurring woody species: Quercus fusiformis, Diospyros texana, Prosopis glandulosa, and Juniperus ashei. We examined hypothesized mechanisms that could promote these species diverse mortality patterns using postdrought measurements on surviving trees coupled to retrospective process modelling. The species exhibited a wide range of gas exchange responses, hydraulic strategies, and mortality rates. Multiple proposed indices of mortality mechanisms were inconsistent with the observed mortality patterns across species, including measures of the degree of iso/anisohydry, photosynthesis, carbohydrate depletion, and hydraulic safety margins. Large losses of spring and summer whole-tree conductance (driven by belowground losses of conductance) and shallower rooting depths were associated with species that exhibited greater mortality. Based on this retrospective analysis, we suggest that species more vulnerable to drought were more likely to have succumbed to hydraulic failure belowground.

Vegetation-zonation patterns across a temperate mountain cloud forest ecotone are not explained by variation in hydraulic functioning or water relations

Many studies have demonstrated linkages between the occurrence of fog and ecophysiological functioning in cloud forests, but few have investigated hydraulic functioning as a determining factor that explains sharp changes in vegetation. The objective of this study was to compare the plant water status during cloud-immersed and non-immersed conditions and hydraulic vulnerability in branches and roots of species across a temperate, mountain fog ecotone. Because cloud forests are often dark, cool and very moist, we expected cloud forest species to have less drought-tolerant characteristics (i.e., lower Pe and P50-the pressures required to induce a 12 and 50% loss in hydraulic conductivity, respectively) relative to non-cloud forest species in adjacent (lower elevation) forests. Additionally, due to the ability of cloud forest species to absorb cloud-fog water, we predicted greater improvements in hydraulic functioning during fog in cloud forest species relative to non-cloud forest species. Across the cloud forest ecotone, most species measured were very resistant to losses in conductivity with branch P50 values from -4.5 to -6.0u2005MPa, hydraulic safety margins (Ψmin - P50) >1.5u2005MPa and low calculated hydraulic conductivity losses. Roots had greater vulnerabilities, with P50 values ranging from -1.4 to -2.5u2005MPa, leading to greater predicted losses in conductivity (∼20%). Calculated values suggested strong losses of midday leaf hydraulic conductance in three of the four species, supporting the hydraulic segmentation hypothesis. In both cloud forest and hardwood species, Ψs were greater on foggy days than sunny days, demonstrating the importance of fog periods to plant water balance across fog regimes. Thus, frequent fog did not result in systemic changes in hydraulic functioning or vulnerability to embolism across our temperate cloud forest ecotone. Finally, roots functioned with lower hydraulic conductivity than branches, suggesting that they may serve as more sensitive indicators of hydraulic functioning in these mesic, foggy ecosystems.

Leaf hydraulic parameters are more plastic in species that experience a wider range of leaf water potentials

Many plant species experience large differences in soil moisture availability within a season, potentially leading to a wide range of leaf water potentials (ΨLEAF). In order to decrease the risk of leaf dehydration, among species, there is a continuum ranging from strict control (isohydry) to little control (anisohydry) of minimum ΨLEAF. In central Texas USA, species are exposed to a range of soil moisture from wet springs to hot, dry summers. There are diverging water management strategies among the four dominant woody species in this system; two of these species are more isohydric (Prosopis glandulosa, Quercus fusiformis) while two others are more anisohydric (Diospyros texana, Juniperus asheii). To maintain leaf turgor and photosynthesis during periods of limited soil moisture, anisohydric species may adjust leaf hydraulic parameters more than isohydric species. To test this hypothesis, we quantified iso/anisohydry from 3xa0years of ΨLEAF predawn and midday measurements, and we measured the changes in turgor loss points (ΨTLP), osmotic potential at full hydration (Ψπ₁₀₀), and resistance to leaf hydraulic dysfunction (leaf P₅₀) throughout the spring and summer of 2016. Diospyros and Juniperus experienced more negative ΨLEAF and adjusted ΨTLP and Ψπ₁₀₀ in response to both drying soils during the summer also in response to rainfall events during September. In contrast, the more isohydric species (Quercus and Prosopis) did not appear to adjust ΨTLP or Ψπ₁₀₀ in response to soil moisture. The more anisohydric species also adjusted leaf P₅₀ during periods of reduced soil moisture. Our results suggest that species that experience wider ranges of ΨLEAF have a greater ability to alter leaf hydraulic properties. This provides insight on how species with different strategies for water potential regulation may modify properties to mitigate drought effects in the future. A plain language summary is available for this article.

Managing Watersheds for Ecosystem Services in the Steepland Neotropics

Este informe sintetiza los resultados de la conferencia Manejo de cuencas hidrograficas para la provision de servicios ambientales en paisajes modificados del neotropico e incluye investigaciones y practicas recientes relacionadas con la gestion de las cuencas hidrograficas de la region. Proporciona una comprension biofisica de la funcion del ecosistema para los usos clave del suelo en el area, resume los servicios ecosistemicos, aborda las consecuencias del cambio climatico y del uso del suelo y proporciona las bases socioeconomicas de los servicios ecosistemicos y los avances en la region. El informe presenta un plan de accion para mejorar la gestion de las cuencas hidrograficas y ofrece casos practicos seleccionados para ilustrar con ejemplos donde se estan realizando avances.This publication represents a synthesis of themes discussed in the following conference Watershed Management for Ecosystem Services in Human Dominated Landscapes of the Neotropics and includes recent research and practices related to watershed management in the region. It provides a biophysical understanding of ecosystem function for key land uses in the area, summarizes ecosystem services, addresses the implications of climate and land use change, and provides socio-economic foundations of ecosystem services and advances in the region. The report presents a road map for improving watershed management and provides selected case studies to illustrate examples of where advances are being made.

Why size matters: the interactive influences of tree diameter distribution and sap flow parameters on upscaled transpiration

In stands with a broad range of diameters, a small number of very large trees can disproportionately influence stand basal area and transpiration (Et). Sap flow-based Et estimates may be particularly sensitive to large trees due to nonlinear relationships between tree-level water use (Q) and tree diameter at breast height (DBH). Because Q is typically predicted on the basis of DBH and sap flow rates measured in a subset of trees and then summed to obtain Et, we assessed the relative importance of DBH and sap flow variables (sap velocity, Vs, and sapwood depth, Rs) in determining the magnitude of Et and its dependence on large trees in a tropical montane forest ecosystem. Specifically, we developed a data-driven simulation framework to vary the relationship between DBH and Vs and stand DBH distribution and then calculate Q, Et and the proportion of Et contributed by the largest tree in each stand. Our results demonstrate that variation in how Rs is determined in the largest trees can alter estimates up to 26% of Et while variation in how Vs is determined can vary results by up to 132%. Taken together, these results highlight a great need to expand our understanding of water transport in large trees as this hinders our ability to predict water fluxes accurately from stand to catchment scales.

Are Northeastern U.S. forests vulnerable to extreme drought

In the Northeastern U.S., drought is expected to increase in frequency over the next century, and therefore, the responses of trees to drought are important to understand. There is recent debate about whether land-use change or moisture availability is the primary driver of changes in forest species composition in this region. Some argue that fire suppression from the early twentieth century to present has resulted in an increase in shade-tolerant and pyrophobic tree species that are drought intolerant, while others suggest precipitation variability as a major driver of species composition. From this debate, an emerging hypothesis is that mesophication and increases in the abundance of mesophytic genera (e.g., Acer, Betula, and Fagus) resulted in forests that are more vulnerable to drought. This review examines the published literature and factors that contribute to drought vulnerability of Northeastern U.S. forests. We assessed two key concepts related to drought vulnerability, including drought tolerance (ability to survive drought) and sensitivity (short-term responses to drought), with a focus on Northeastern U.S. species. We assessed drought-tolerance classifications for species, which revealed both consistencies and inconsistencies, as well as contradictions when compared to actual observations, such as higher mortality for drought-tolerant species. Related to drought sensitivity, recent work has focused on isohydric/anisohydric regulation of leaf water potential. However, based on the review of the literature, we conclude that drought sensitivity should be viewed in terms of multiple variables, including leaf abscission, stomatal sensitivity, turgor pressure, and dynamics of non-structural carbohydrates. Genera considered drought sensitive (e.g., Acer, Betula, and Liriodendron) may actually be less prone to drought-induced mortality and dieback than previously considered because stomatal regulation and leaf abscission in these species are effective at preventing water potential from reaching critical thresholds during extreme drought. Independent of drought-tolerance classification, trees are prone to dieback and mortality when additional stressors are involved such as insect defoliation, calcium and magnesium deficiency, nitrogen saturation, and freeze-thaw events. Overall, our literature review shows that multiple traits associated with drought sensitivity and tolerance are important as species may rely on different mechanisms to prevent hydraulic failure and depleted carbon reserves that may lead to mortality.