Rainforest conversion to woody croplands impacts the carbon cycle via ecophysiological processes such as photosynthesis and autotrophic respiration. Changes in the carbon cycle associated with land-use change can be estimated through Land Surface Models (LSMs). The accuracy of carbon flux estimation in carbon fluxes associated with land-use change has been attributed to uncertainties in the model parameters affecting photosynthetic activity, which is a function of both carboxylation capacity (Vcmax) and electron transport capacity (Jmax). In order to reduce such uncertainties for common tropical woody crops and trees, in this study we measured Vcmax25 (Vcmax standardized to 25 °C), Jmax25 (Jmax standardized to 25 °C) and light-saturated photosynthetic capacity (Amax) of Elaeis guineensis Jacq. (oil palm), Hevea brasiliensis (rubber tree), and two native tree species, Eusideroxylon zwageri and Alstonia scholaris, in a converted landscape in Jambi province (Sumatra, Indonesia) at smallholder plantations. We considered three plantations; a monoculture rubber, a monoculture oil palm, and an agroforestry system (jungle rubber plantation), where rubber trees coexist with some native trees. We performed measurements on leaves at the lower part of the canopy, and used a scaling method based on exponential function to scale up photosynthetic capacity related traits to the top of the canopy. At the lower part of the canopy, we found (i) high Vcmax25 values for H. brasiliensis from monoculture rubber plantation and jungle rubber plantation that was linked to a high area-based leaf nitrogen content, and (ii) low value of Amax for E. guineensis from oil palm plantation that was due to a low value of Vcmax25 and a high value of dark respiration. At the top of the canopy, Amax varied much more than Vcmax25 among different land-use types. We found that photosynthetic capacity declined fastest from the top to the lower part of the canopy in oil palm plantations. We demonstrate that photosynthetic capacity related traits measured at the lower part of the canopy can be successfully scaled up to the top of the canopy. We thus provide helpful new data that can be used to constrain LSMs that simulate land-use change related to rubber and oil palm expansion.

Abstract Diurnal branch movements in woody plants have only recently been described in detail. While previously only vegetative and reproductive structures have been known to move on hourly timescales, imaging technologies such as terrestrial laser scanning and near‐surface repeat digital photography provide a means of remotely monitoring plant movements at high enough temporal and spatial resolution to capture rhythmic movements of woody material. Virtually, nothing is known about the range of species and ecosystems in which woody movements might occur or what causes these movements. We report that diurnal woody branch movements occur in a number of tree and shrub species across a broad range of abiotic conditions. We examined detailed branch movements in one species, creosote (Larrea tridentata), and found that branch movements were highly correlated with humidity, air temperature, vapor pressure deficit, and stem water potential: all factors related to plant water status. We also found that live and dead branch movements were distinct in the timing of their movements and in the abiotic conditions with which they were most correlated. Changes in dead branch position were most correlated with humidity, with these movements consistently lagging 1–2 h behind changes in humidity. Live branch movements were also highly correlated with vapor pressure deficit and humidity but went from lagging 1–2 h behind changes in these abiotic conditions in summer to being nearly in sync in winter. We believe that this is the first study that (1) documents diurnal branch movements in creosote, (2) differentiates between the movements of live and dead branches, and (3) relates environmental data to these movements. We hope these findings encourage other researchers to more closely examine imagery from their sites for evidence of branch movements, which may provide deeper insights into water and solute movements in plants and physiological responses to water stress.

Plants close their stomata during drought to avoid excessive water loss, but species differ in respect to the drought severity at which stomata close. The stomatal closure point is related to xylem anatomy and vulnerability to embolism, but it also has implications for phloem transport and possibly phloem anatomy to allow sugar transport at low water potentials. Desiccation-tolerant plants that close their stomata at severe drought should have smaller xylem conduits and/or fewer and smaller interconduit pits to reduce vulnerability to embolism but more phloem tissue and larger phloem conduits compared with plants that avoid desiccation. These anatomical differences could be expected to increase in response to long-term reduction in precipitation. To test these hypotheses, we used tridimensional synchroton X-ray microtomograph and light microscope imaging of combined xylem and phloem tissues of 2 coniferous species: one-seed juniper (Juniperus monosperma) and piñon pine (Pinus edulis) subjected to precipitation manipulation treatments. These species show different xylem vulnerability to embolism, contrasting desiccation tolerance, and stomatal closure points. Our results support the hypothesis that desiccation tolerant plants require higher phloem transport capacity than desiccation avoiding plants, but this can be gained through various anatomical adaptations in addition to changing conduit or tissue size.

Tree transpiration depends on biotic and abiotic factors that might change in the future, including precipitation and soil moisture status. Although short-term sap flux responses to soil moisture and evaporative demand have been the subject of attention before, the relative sensitivity of sap flux to these two factors under long-term changes in soil moisture conditions has rarely been determined experimentally. We tested how long-term artificial change in soil moisture affects the sensitivity of tree-level sap flux to daily atmospheric vapor pressure deficit (VPD) and soil moisture variations, and the generality of these effects across forest types and environments using four manipulative sites in mature forests. Exposure to relatively long-term (two to six years) soil moisture reduction decreases tree sap flux sensitivity to daily VPD and relative extractable water (REW) variations, leading to lower sap flux even under high soil moisture and optimal VPD. Inversely, trees subjected to long-term irrigation showed a significant increase in their sensitivity to daily VPD and REW, but only at the most water-limited site. The ratio between the relative change in soil moisture manipulation and the relative change in sap flux sensitivity to VPD and REW variations was similar across sites suggesting common adjustment mechanisms to long-term soil moisture status across environments for evergreen tree species. Overall, our results show that long-term changes in soil water availability, and subsequent adjustments to these novel conditions, could play a critical and increasingly important role in controlling forest water use in the future.

Hydraulic architecture imposes a fundamental control on water transport, underpinning plant productivity, and survival. The extent to which hydraulic architecture of mature trees acclimates to chronic drought is poorly understood, limiting accuracy in predictions of forest responses to future droughts. We measured seasonal shoot hydraulic performance for multiple years to assess xylem acclimation in mature pinon (Pinus edulis) and juniper (Juniperus monosperma) after 3+years of precipitation manipulation. Our treatments consisted of water addition (+20% ambient precipitation), partial precipitation-exclusion (-45% ambient precipitation), and exclusion-structure control. Supplemental watering elevated leaf water potential, sapwood-area specific hydraulic conductivity, and leaf-area specific hydraulic conductivity relative to precipitation exclusion. Shifts in allocation of leaf area to sapwood area enhanced differences between irrigated and droughted K-L in pinon but not juniper. Pinon and juniper achieved similar K-L under ambient conditions, but juniper matched or outperformed pinon in all physiological measurements under both increased and decreased precipitation treatments. Embolism vulnerability and xylem anatomy were unaffected by treatments in either species. Absence of significant acclimation combined with inferior performance for both hydraulic transport and safety suggests pinon has greater risk of local extirpation if aridity increases as predicted in the southwestern USA. The unknown capacity of mature trees to acclimate their hydraulic architecture to climate forcing is a source of uncertainty in predicting the response of long-lived individuals to progressive climate change. We used a multi-year in situ precipitation-manipulation experiment to study hydraulic performance of mature pinon and juniper. When subjected to supplemental watering or experimental drought, neither species demonstrated significant adjustments in either hydraulic transport safety or efficiency. We conclude that the hydraulic architecture of mature trees in pinon-juniper woodlands is unlikely to acclimate to predicted climate changes, leaving trees more susceptible to the effects of increased temperature and water stress.

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.

Remotely sensed microwave observations of vegetation optical depth (VOD) have been widely used for examining vegetation responses to climate. Nevertheless, the relative impacts of phenological changes in leaf biomass and water stress on VOD have not been explicitly disentangled. In particular, determining whether leaf water potential ( ψ L ) affects VOD may allow these data sets as a constraint for plant hydraulic models. Here we test the sensitivity of VOD to variations in ψ L and present a conceptual framework that relates VOD to ψ L and total biomass including leaves, whose dynamics are measured through leaf area index, and woody components. We used measurements of ψ L from three sites across the US—a mixed deciduous forests in Indiana and Missouri and a piñon‐juniper woodland in New Mexico—to validate the conceptual model. The temporal dynamics of X‐band VOD were similar to those of the VOD signal estimated from the new conceptual model with observed ψ L ( R 2 = 0.6–0.8). At the global scale, accounting for a combination of biomass and estimated ψ L (based on satellite surface soil moisture data) increased correlations with VOD by ~ 15% and 30% compared to biomass and water potential, respectively. In wetter regions with denser and taller canopy heights, VOD has a higher correlation with leaf area index than with water stress and vice versa in drier regions. Our results demonstrate that variations in both phenology and ψ L must be considered to accurately interpret the dynamics of VOD observations for ecological applications.

Summary Ectotherms such as lizards are expected to alter their behaviour and microhabitat use and experience population declines in response to rising temperatures. But the role of changing rainfall patterns on lizard behaviour and microhabitat use is not understood. We used a 5‐year rainfall manipulation experiment in a piñon pine‐juniper woodland in central New Mexico to study how a lizard species' microhabitat use varies in four different rainfall treatments. We examined ground temperatures in the sun and shade, and daily rainfall, within each treatment, during lizard activity periods, to address how lizards used sun or shade microhabitats on a daily basis. Our small‐scale rainfall and temperature measurements indicate that rainfall, not temperatures, predict lizard microhabitat use. Lizards showed a strong preference for shaded microhabitats during dry periods and used sunny microhabitats following rainfall events, regardless of treatment type. Lizards remained active during dry periods, foraging in the shade of trees. Our study shows that rainfall can influence lizard microhabitat use more than temperature in a piñon pine‐juniper woodland and the trees provide important refugia. The loss of piñon pine and juniper trees from prolonged drought threatens to limit the amount of shade available to lizards in the future.

Annual streamflows have decreased across mountain watersheds in the Pacific Northwest of the United States over the last similar to 70years; however, in some watersheds, observed annual flows have increased. Physically based models are useful tools to reveal the combined effects of climate and vegetation on long-term water balances by explicitly simulating the internal watershed hydrological fluxes that affect discharge. We used the physically based Simultaneous Heat and Water (SHAW) model to simulate the inter-annual hydrological dynamics of a 4km(2) watershed in northern Idaho. The model simulates seasonal and annual water balance components including evaporation, transpiration, storage changes, deep drainage, and trends in streamflow. Independent measurements were used to parameterize the model, including forest transpiration, stomatal feedback to vapour pressure, forest properties (height, leaf area index, and biomass), soil properties, soil moisture, snow depth, and snow water equivalent. No calibrations were applied to fit the simulated streamflow to observations. The model reasonably simulated the annual runoff variations during the evaluation period from water year 2004 to 2009, which verified the ability of SHAW to simulate the water budget in this small watershed. The simulations indicated that inter-annual variations in streamflow were driven by variations in precipitation and soil water storage. One key parameterization issue was leaf area index, which strongly influenced interception across the catchment. This approach appears promising to help elucidate the mechanisms responsible for hydrological trends and variations resulting from climate and vegetation changes on small watersheds in the region. Copyright (c) 2015 John Wiley & Sons, Ltd.

Drought-induced forest mortality is an increasing global problem with wide-ranging consequences, yet mortality mechanisms remain poorly understood. Depletion of non-structural carbohydrate (NSC) stores has been implicated as an important mechanism in drought-induced mortality, but experimental field tests are rare. We used an ecosystem-scale precipitation manipulation experiment to evaluate leaf and twig NSC dynamics of two co-occurring conifers that differ in patterns of stomatal regulation of water loss and recent mortality: the relatively desiccation-avoiding piñon pine (Pinus edulis) and the relatively desiccation-tolerant one-seed juniper (Juniperus monosperma). Piñon pine experienced 72% mortality after 13-25 months of experimental drought and juniper experienced 20% mortality after 32-47 months. Juniper maintained three times more NSC in the foliage than twigs, and converted NSC to glucose and fructose under drought, consistent with osmoregulation requirements to maintain higher stomatal conductance during drought than piñon. Despite these species differences, experimental drought caused decreased leaf starch content in dying trees of both species (P < 0.001). Average dry-season leaf starch content was also a good predictor of drought-survival time for both species (R(2)  = 0.93). These results, along with observations of drought-induced reductions to photosynthesis and growth, support carbon limitation as an important process during mortality of these two conifer species.

Pinon pine-juniper (Pinus edulis-Juniperus monosperma) woodlands constitute a large proportion of land area in the southwestern United States and have experienced widespread vegetation mortality during regional drought events over the past century. Pinon pines have been especially affected by these events, and drought severity is predicted to increase in this region in the future. Based on research that suggests winter climate may influence growing season productivity in semiarid ecosystems, we evaluated the potential for small changes in average winter climate to affect spring growing season conditions in pinon-juniper woodlands, New Mexico, USA. We developed a low-dimensional ecohydrological model of pinon-juniper woodland ecosystems on moderate slopes (5%) and on steep slopes (25%) and simulated the responses of ecosystem water availability, surface conditions, and water and carbon flux dynamics to a climate change scenario of increased temperature and decreased winter precipitation. The climate change scenario reduced average winter snowcover, decreased surface albedo, increased net radiation, and altered the timing of spring evaporation (E) towards earlier dates. Moderate slope pinon and juniper trees experienced small reductions in transpiration (Tr) and carbon assimilation (A), and those on steep slopes experienced small but relatively larger reductions in Tr and A, as well as higher increases in soil moisture (0) variance and E variance. As a result of climate change, the peak of spring Tr occurred on average 6 days earlier on moderate slopes and 10 days earlier on steep slopes, the timing of A shifted towards earlier March dates, and A was reduced during April and May. Steep slope pinon pines experienced greater proportional reductions in Tr and A than junipers. Our results suggests that winter climate change will promote an earlier growing season in pinon-juniper woodlands, will increase daily variance in 9 and E during spring, and will produce slight reductions in Am woodlands with steep slopes and a large proportion of pinon pines. In a more arid future climate, a shift towards lower soil moisture availability and carbon assimilation in April and May may intensify the effects of early summer drought events for pinon-juniper woodlands, thus exacerbating the impacts of larger changes in climate dynamics. Published by Elsevier B.V.

Plant hydraulic conductance (ks) is a critical control on whole-plant water use and carbon uptake and, during drought, influences whether plants survive or die. To assess long-term physiological and hydraulic responses of mature trees to water availability, we manipulated ecosystem-scale water availability from 2007 to 2013 in a piñon pine (Pinus edulis) and juniper (Juniperus monosperma) woodland. We examined the relationship between ks and subsequent mortality using more than 5 years of physiological observations, and the subsequent impact of reduced hydraulic function and mortality on total woody canopy transpiration (EC) and conductance (GC). For both species, we observed significant reductions in plant transpiration (E) and ks under experimentally imposed drought. Conversely, supplemental water additions increased E and ks in both species. Interestingly, both species exhibited similar declines in ks under the imposed drought conditions, despite their differing stomatal responses and mortality patterns during drought. Reduced whole-plant ks also reduced carbon assimilation in both species, as leaf-level stomatal conductance (gs) and net photosynthesis (An) declined strongly with decreasing ks. Finally, we observed that chronically low whole-plant ks was associated with greater canopy dieback and mortality for both piñon and juniper and that subsequent reductions in woody canopy biomass due to mortality had a significant impact on both daily and annual canopy EC and GC. Our data indicate that significant reductions in ks precede drought-related tree mortality events in this system, and the consequence is a significant reduction in canopy gas exchange and carbon fixation. Our results suggest that reductions in productivity and woody plant cover in piñon–juniper woodlands can be expected due to reduced plant hydraulic conductance and increased mortality of both piñon pine and juniper under anticipated future conditions of more frequent and persistent regional drought in the southwestern United States.

Interspecific variation in stomatal conductance (GS) and transpiration (EL) has been documented in stands of co-occurring species, and this variation has been observed to differ with tree size and canopy height increase. In this study, we present data that examine fluctuations in canopy gas exchange across co-occurring species and varying canopy heights for three montane forest chronosequences located in an inland Pacific Northwest mixed-conifer forest. With the exception of Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco), we observed consistent declines in canopy EL and GS with increasing height for the majority of species examined in our 2-year study. Along with declines in canopy GS, we observed decreases in leaf-specific hydraulic conductance (KL) across species as canopy height increased. Seasonally, we observed declines in canopy GS during warmer and dryer summer months of both years. These decreases in GS were significant (up to 50%) and suggest that carbon assimilation in trees was limited during dryer months due to a combination of high evaporative demand and reduced soil H2O availability. Such reductions in GS during periods of increased plant water stress suggest that forest productivity in the inland Pacific Northwest may be impacted negatively if future climate predictions of increasing growing-season water stress are realized.

Fundamental drivers of ecosystem processes such as temperature and precipitation are rapidly changing and creating novel environmental conditions. Forest landscape models ( FLM ) are used by managers and policy‐makers to make projections of future ecosystem dynamics under alternative management or policy options, but the links between the fundamental drivers and projected responses are weak and indirect, limiting their reliability for projecting the impacts of climate change. We developed and tested a relatively mechanistic method to simulate the effects of changing precipitation on species competition within the LANDIS ‐ II FLM . Using data from a field precipitation manipulation experiment in a piñon pine ( Pinus edulis ) and juniper ( Juniperus monosperma ) ecosystem in New Mexico ( USA ), we calibrated our model to measurements from ambient control plots and tested predictions under the drought and irrigation treatments against empirical measurements. The model successfully predicted behavior of physiological variables under the treatments. Discrepancies between model output and empirical data occurred when the monthly time step of the model failed to capture the short‐term dynamics of the ecosystem as recorded by instantaneous field measurements. We applied the model to heuristically assess the effect of alternative climate scenarios on the piñon–juniper ecosystem and found that warmer and drier climate reduced productivity and increased the risk of drought‐induced mortality, especially for piñon. We concluded that the direct links between fundamental drivers and growth rates in our model hold great promise to improve our understanding of ecosystem processes under climate change and improve management decisions because of its greater reliance on first principles.

A semi-mechanistic forest growth model, 3-PG (Physiological Principles Predicting Growth), was extended to calculate C-13 in tree rings. The C-13 estimates were based on the model's existing description of carbon assimilation and canopy conductance. The model was tested in two approximate to 80-year-old natural stands of Abies grandis (grand fir) in northern Idaho. We used as many independent measurements as possible to parameterize the model. Measured parameters included quantum yield, specific leaf area, soil water content and litterfall rate. Predictions were compared with measurements of transpiration by sap flux, stem biomass, tree diameter growth, leaf area index and C-13. Sensitivity analysis showed that the model's predictions of C-13 were sensitive to key parameters controlling carbon assimilation and canopy conductance, which would have allowed it to fail had the model been parameterized or programmed incorrectly. Instead, the simulated C-13 of tree rings was no different from measurements (P>0.05). The C-13 submodel provides a convenient means of constraining parameter space and avoiding model artefacts. This C-13 test may be applied to any forest growth model that includes realistic simulations of carbon assimilation and transpiration.