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1. Occult precipitation events (fog, dew, light-rain) can alter plant water and nutritional status, both directly through the aerial uptake of surface water and nutrients, and indirectly via redistribution of atmospheric resources to the soil. However, current frameworks that explain niche segregation, species interactions, and coexistence still consider that ground-rooted plants obtain resources almost exclusively via root absorption from soil.2. Here, we expand the plant hydrological niches model to incorporate both soil and atmospheric resource-axes, thus providing a more complete picture of how ground-rooted terrestrial plants obtain, remobilize, share, and compete for water and soluble nutrients.3. First, we describe how plants with different water acquisition strategies access directly or indirectly atmospheric resources. Then, we discuss how the use of such resources may promote spatiotemporal niche segregation, contributing to shape species distribution and abundance within plant communities. We illustrate this argument with examples from arid, mesic, and wet vegetation types. Finally, we examine how climate and land-use changes may influence plant hydrological niches, potentially altering community structure.4. Synthesis: Understanding how available atmospheric resources influences niche segregation in plant communities is a crucial step towards better predictions of species responses (e.g., changes in distribution, abundance and interactions) to climate change.

Summary-Producing and maintaining flowers is essential for reproduction in most angiosperms, underpinning population persistence and speciation. Although the physiological costs of flowers often oppose pollinator selection, these physiological costs have rarely been quantified.-We measured a suite of physiological traits quantifying the water and carbon costs and drought tolerance on flowers and leaves of over 100 phylogenetically diverse species, including water and dry mass contents, minimum epidermal conductance to water vapor (gmin), vein density, and dry mass per area.-Although there was substantial variation among species, flowers had significantly highergminand water content per unit area than leaves, but significantly lower vein density and dry mass per area than leaves. Both leaves and flowers exhibited similarly strong scaling between dry mass investment and water content.-The highergminof flowers offset their higher water content, suggesting that flowers may desiccate more rapidly than leaves during drought. The coordination between dry mass and water investment suggests that flowers rely on a hydrostatic skeleton to remain upright rather than on a carbon-based skeleton. For short-lived structures like flowers, water may be relatively cheaper than carbon, particularly given the relatively high amount of water loss per unit of carbon synthesized in photosynthesis.

Understanding climate change impacts on the Cape Floristic Region requires improved knowledge of plant physiological responses to the environment. Studies examining physiological responses of mountain fynbos have consisted of campaign-based measurements, capturing snapshots of plant water relations and photosynthesis. We examine conclusions drawn from prior studies by tracking in situ physiological responses of three species, representing dominant growth forms (proteoid, ericoid, restioid), over 2 years using miniature continuous sap flow technology, long-term observations of leaf/culm water potential and gas exchange, and xylem vulnerability to embolism. We observed considerable inter-specific variation in the timing and extent of seasonal declines in productivity. Shallow-rooted Erica monsoniana exhibited steep within-season declines in sap flow and water potentials, and pronounced inter-annual variability in total daily sap flux (Js). Protea repens showed steady reductions in Js across both years, despite deeper roots and less negative water potentials. Cannomois congesta—a shallow-rooted restioid—was least negatively impacted. Following rehydrating rain at the end of summer, gas exchange recovery was lower in the drier year compared with the normal year, but did not differ between species. Loss of function in the drier year was partially accounted for by loss of xylem transport capacity in Erica and Cannomois, but not Protea. Hitherto unseen water use patterns, including inter-annual variability of gas exchange associated with contrasting water uptake properties, reveal that species use different mechanisms to cope with summer dry periods. Revealing physiological responses of key growth forms enhances predictions of plant function within mountain fynbos under future conditions.

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.

Dew is an important water resource for plants in most deserts. The mechanism that allows desert plants to use dew water was studied using an isotopic water tracer approach. Most plants use water directly from the soil; the roots transfer the water to the rest of the plant, where it is required for all metabolic functions. However, many plants can also take up water into their leaves and stems. Examining the dew water uptake pathways in desert plants can lend insight on another all water-use pathways examination. We determined where and how dew water enters plants in the water limited Negev desert. Highly depleted isotopic water was sprayed on three different dominant plant species of the Negev desert—Artemesia sieberi, Salsola inermis and Haloxylon scoparium—and its entry into the plant was followed. Water was sprayed onto the soil only, or on the leaves/stems only (with soil covered to prevent water entry via root uptake). Thereafter, the isotopic composition of water in the roots and stems were measured at various time points. The results show that each plant species used the dew water to a different extent, and we obtained evidence of foliar uptake capacity of dew water that varied depending on the microenvironmental conditions. A. sieberi took up the greatest amount of dew water through both stems and roots, S. inermis took up dew water mainly from the roots, and H. scoparium showed the least dew capture overall.

Experimental drought has been shown to delay the development of the root microbiome and increase the relative abundance of Actinobacteria, however, the generalizability of these findings to natural systems or other diverse plant hosts remains unknown. Bacterial cell wall thickness and growth morphology (e.g., filamentous or unicellular) have been proposed as traits that may mediate bacterial responses to environmental drivers. Leveraging a natural gradient of water-availability across the coast redwood (Sequoia sempervirens) range, we tested three hypotheses: (a) that site-specific water-availability is an important predictor of bacterial community composition for redwood roots and rhizosphere soils; (b) that there is relative enrichment of Actinobacteria and other monoderm bacterial groups within the redwood microbiome in response to drier conditions; and (c) that bacterial growth morphology is an important predictor of bacteria response to water-availability, where filamentous taxa will become more dominant at drier sites compared to unicellular bacteria. We find that both α- and β-diversity of redwood bacterial communities is partially explained by water-availability and that Actinobacterial enrichment is a conserved response of land plants to water-deficit. Further, we highlight how the trend of Actinobacterial enrichment in the redwood system is largely driven by the Actinomycetales. We propose bacterial growth morphology (filamentous vs. unicellular) as an additional mechanism behind the increase in Actinomycetales with increasing aridity. A trait-based approach including cell-wall thickness and growth morphology may explain the distribution of bacterial taxa across environmental gradients and help to predict patterns of bacterial community composition for a wide range of host plants.

Amazonian droughts are increasing in frequency and severity. However, little is known about how this may influence species-specific vulnerability to drought across different ecosystem types. We measured 16 functional traits for 16 congeneric species from six families and eight genera restricted to floodplain, swamp, white-sand or plateau forests of Central Amazonia. We investigated whether habitat distributions can be explained by species hydraulic strategies, and if habitat specialists differ in their vulnerability to embolism that would make water transport difficult during drought periods. We found strong functional differences among species. Nonflooded species had higher wood specific gravity and lower stomatal density, whereas flooded species had wider vessels, and higher leaf and xylem hydraulic conductivity. The P50 values (water potential at 50% loss of hydraulic conductivity) of nonflooded species were significantly more negative than flooded species. However, we found no differences in hydraulic safety margin among species, suggesting that all trees may be equally likely to experience hydraulic failure during severe droughts. Water availability imposes a strong selection leading to differentiation of plant hydraulic strategies among species and may underlie patterns of adaptive radiation in many tropical tree genera. Our results have important implications for modeling species distribution and resilience under future climate scenarios.

Root-associated fungal communities modify the climatic niches and even the competitive ability of their hosts, yet how the different components of the root microbiome are modified by habitat loss remains a key knowledge gap. Using principles of landscape ecology, we tested how free-living versus host-associated microbes differ in their response to landscape heterogeneity. Further, we explore how compartmentalisation of microbes into specialised root structures filters for key fungal symbionts. Our study demonstrates that free-living fungal community structure correlates with landscape heterogeneity, but that host-associated fungal communities depart from these patterns. Specifically, biotic filtering in roots, especially via compartmentalisation within specialised root structures, decouples the biogeographic patterns of host-associated fungal communities from the soil community. In this way, even as habitat loss and fragmentation threaten fungal diversity in the soils, plant hosts exert biotic controls to ensure associations with critical mutualists, helping to preserve the root mycobiome.

Climate warming in recent decades has negatively impacted forest health in the western United States. Here, we report on potential early warning signals (EWS) for drought-related mortality derived from measurements of tree-ring growth (ring width index; RWI) and carbon isotope discrimination (∆13 C), primarily focused on ponderosa pine (Pinus ponderosa). Sampling was conducted in the southern Sierra Nevada Mountains, near the epicenter of drought severity and mortality associated with the 2012-2015 California drought and concurrent outbreak of western pine beetle (Dendroctonus brevicomis). At this site, we found that widespread mortality was presaged by five decades of increasing sensitivity (i.e., increased explained variation) of both tree growth and ∆13 C to Palmer Drought Severity Index (PDSI). We hypothesized that increasing sensitivity of tree growth and ∆13 C to hydroclimate constitute EWS that indicate an increased likelihood of widespread forest mortality caused by direct and indirect effects of drought. We then tested these EWS in additional ponderosa pine-dominated forests that experienced varying mortality rates associated with the same California drought event. In general, drier sites showed increasing sensitivity of RWI to PDSI over the last century, as well as higher mortality following the California drought event compared to wetter sites. Two sites displayed evidence that thinning or fire events that reduced stand basal area effectively reversed the trend of increasing hydroclimate sensitivity. These comparisons indicate that reducing competition for soil water and/or decreasing bark beetle host tree density via forest management-particularly in drier regions-may buffer these forests against drought stress and associated mortality risk. EWS such as these could provide land managers more time to mitigate the extent or severity of forest mortality in advance of droughts. Substantial efforts at deploying additional dendrochronological research in concert with remote sensing and forest modeling will aid in forecasting of forest responses to continued climate warming.

Klein (1) challenges two interpretations in Niespolo et al. (2). Regarding his first point, we maintain that Ysterfontein 1 (YFT1) does provide the oldest known example of full coastal adaptation as indicated by the presence of shell middens (cf. ref. 3). Klein inaccurately characterizes the age of the deepest shell midden layers at Klasies River Main (KRM) given in ref. 4. While other workers identify even the oldest light brown sand (LBS) layer at KRM as similar in age or younger than the deepest deposits at YFT1 (cf. refs. 5 and 6), the reference cited by Klein (4) and subsequent dating efforts show that the shell midden lenses in beach-sand deposits within the LBS layer likely postdate the marine isotope stage 5e sea level highstand (∼120 ka) as does the midden at YFT1. However, existing dates suggest that KRM shell middens may, in fact, be younger than YFT1. Layer LBS has been directly dated using multiple techniques that converge on an age of ∼104 ka (5), notably younger than the 95% confidence interval for YFT1 deposition (2). However, complicating matters, a capping speleothem with a U-series age of ∼110 ka is interpreted as a minimum age of the LBS layer at KRM (5). Thus, layer LBS yields conflicting dates and its true age remains uncertain. The next oldest layer at KRM hosting midden deposits (layer shell and sand [SAS]) is definitively younger than YFT1 (5, 7). Thus, there is no basis for the assertion that KRM middens are older than those at YFT1, and we stand by our claim that YFT1 provides the oldest currently known example of systematic coastal resource exploitation.

Plants are critical mediators of terrestrial mass and energy fluxes, and their structural and functional traits have profound impacts on local and global climate, biogeochemistry, biodiversity, and hydrology. Yet, Earth System Models (ESMs), our most powerful tools for predicting the effects of humans on the coupled biosphere-atmosphere system, simplify the incredible diversity of land plants into a handful of coarse categories of "Plant Functional Types" (PFTs) that often fail to capture ecological dynamics such as biome distributions. The inclusion of more realistic functional diversity is a recognized goal for ESMs, yet there is currently no consistent, widely accepted way to add diversity to models, that is, to determine what new PFTs to add and with what data to constrain their parameters. We review approaches to representing plant diversity in ESMs and draw on recent ecological and evolutionary findings to present an evolution-based functional type approach for further disaggregating functional diversity. Specifically, the prevalence of niche conservatism, or the tendency of closely related taxa to retain similar ecological and functional attributes through evolutionary time, reveals that evolutionary relatedness is a powerful framework for summarizing functional similarities and differences among plant types. We advocate that Plant Functional Types based on dominant evolutionary lineages ("Lineage Functional Types") will provide an ecologically defensible, tractable, and scalable framework for representing plant diversity in next-generation ESMs, with the potential to improve parameterization, process representation, and model benchmarking. We highlight how the importance of evolutionary history for plant function can unify the work of disparate fields to improve predictive modeling of the Earth system.

Vulnerability to embolism varies between con-generic species distributed along aridity gradients, yet little is known about intraspecific variation and its drivers. Even less is known about intraspecific variation in tissues other than stems, despite results suggesting that roots, stems and leaves can differ in vulnerability. We hypothesized that intraspecific variation in vulnerability in leaves and stems is adaptive and driven by aridity. We quantified leaf and stem vulnerability of Quercus douglasii using the optical technique. To assess contributions of genetic variation and phenotypic plasticity to within-species variation, we quantified the vulnerability of individuals growing in a common garden, but originating from populations along an aridity gradient, as well as individuals from the same wild populations. Intraspecific variation in water potential at which 50% of total embolism in a tissue is observed (P50 ) was explained mostly by differences between individuals (>66% of total variance) and tissues (16%). There was little between-population variation in leaf/stem P50 in the garden, which was not related to site of origin aridity. Unexpectedly, we observed a positive relationship between wild individual stem P50 and aridity. Although there is no local adaptation and only minor phenotypic plasticity in leaf/stem vulnerability in Q. douglasii, high levels of potentially heritable variation within populations or strong environmental selection could contribute to adaptive responses under future climate change.

Fog and low-lying cloud (fog) play a significant role in the maintenance of ecosystems, from desert to alpine and from coastal to inland systems. Our central thesis is that fog provides ecosystems with critical water and nutrient subsidies, and also delivers pollutants, that often control ecosystem function. Fog is a medium, vector, and connector. In this mini-review, we synthesize recent research advances that reveal the diverse ways that fog shapes ecosystem processes. Crown wetting, elemental deposition, and light scattering and absorption are fundamental mechanisms by which fog has been shown to influence water fluxes, productivity, and decomposition in hyper-arid to ever-wet regions. These impacts are ultimately mediated by the structure and composition of biological systems that allow fog capture and utilization of resource subsidies. Climate change, and changes in land use, ocean circulation, and atmospheric pollution are simultaneously altering the nature of fog itself, and the architecture of the ecosystems adapted to capture it. The coupling between atmosphere and biosphere in fog-enshrouded areas raises new questions about past and future fog-dominated ecosystems, and their maintenance and diversity, in the face of global change.

In the Earth’s tallest tree species, axial variation in conduit size matches theoretical predictions minimizing the accumulation of hydraulic resistance with height growth, within a constraint of maximum conduit diameter. Hydraulic limitations to tree height can be mitigated by widening the conducting elements toward a tree’s base. However, size limits of tracheid and vessel dimensions may constrain this compensation mechanism as the water transport pathway elongates. Moreover, variation in conduit size is poorly described in tall trees even though their long transport paths have high potential for hydraulic resistance. Here, we evaluated whether axial variation in conduit diameter was uniquely structured, or matched theoretical predictions in Sequoia sempervirens, Sequoiadendron giganteum, and Eucalyptus regnans that were 86–105 m tall and exceeded 85% of the maximum height for each species. Across Sequoia and Sequoiadendron, tree top tracheids maintained constant width, whereas tree base tracheids in the outermost ring were 20% wider in taller trees, suggesting maintenance of basipetal conduit widening with height growth. In all trees, the observed widening decreased at a rate per unit path length that fitted well to a power function with an exponent consistent with hydraulic compensation. However, below about 60 m from the tree tops, conduit diameters approached an asymptote beneath the power function, indicating a limit to maximum conduit size. Quantifying the distribution of base-to-top hydraulic resistance suggested that the minimal hydraulic benefit gained with increasingly wider conduits near the tree base may trade off with other factors such as maintaining mechanical strength or reducing fluid volume. We summarize these results into an anatomical model of height growth that includes limits to axial variation in conduit diameter and is supported by many physiological and anatomical observations.

Despite the appeal of the iso/anisohydric framework for classifying plant drought responses, recent studies have shown that such classifications can be strongly affected by a plant's environment. Here, we present measured in situ drought responses to demonstrate that apparent isohydricity can be conflated with environmental conditions that vary over space and time. In particular, we (a) use data from an oak species (Quercus douglasii) during the 2012–2015 extreme drought in California to demonstrate how temporal and spatial variability in the environment can influence plant water potential dynamics, masking the role of traits; (b) explain how these environmental variations might arise from climatic, topographic, and edaphic variability; (c) illustrate, through a “common garden” thought experiment, how existing trait-based or response-based isohydricity metrics can be confounded by these environmental variations, leading to Type-1 (false positive) and Type-2 (false negative) errors; and (d) advocate for the use of model-based approaches for formulating alternate classification schemes. Building on recent insights from greenhouse and vineyard studies, we offer additional evidence across multiple field sites to demonstrate the importance of spatial and temporal drivers of plants' apparent isohydricity. This evidence challenges the use of isohydricity indices, per se, to characterize plant water relations at the global scale. This is the peer reviewed version of the following article: Feng, X, Ackerly, DD, Dawson, TE, et al. Beyond isohydricity: The role of environmental variability in determining plant drought responses. Plant Cell Environ. 2019; 42: 1104– 1111, which has been published in final form at https://doi.org/10.1111/pce.13486. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Versions. This article may not be enhanced, enriched or otherwise transformed into a derivative work, without express permission from Wiley or by statutory rights under applicable legislation. Copyright notices must not be removed, obscured or modified. The article must be linked to Wiley’s version of record on Wiley Online Library and any embedding, framing or otherwise making available the article or pages thereof by third parties from platforms, services and websites other than Wiley Online Library must be prohibited. &nbsp

Modern human behavioral innovations from the Middle Stone Age (MSA) include the earliest indicators of full coastal adaptation evidenced by shell middens, yet many MSA middens remain poorly dated. We apply (230)Th/U burial dating to ostrich eggshells (OES) from Ysterfontein 1 (YFT1, Western Cape, South Africa), a stratified MSA shell midden. (230)Th/U burial ages of YFT1 OES are relatively precise (median ± 2.7%), consistent with other age constraints, and preserve stratigraphic principles. Bayesian age–depth modeling indicates YFT1 was deposited between 119.9 to 113.1 thousand years ago (ka) (95% CI of model ages), and the entire 3.8 m thick midden may have accumulated within ∼2,300 y. Stable carbon, nitrogen, and oxygen isotopes of OES indicate that during occupation the local environment was dominated by C(3) vegetation and was initially significantly wetter than at present but became drier and cooler with time. Integrating archaeological evidence with OES (230)Th/U ages and stable isotopes shows the following: 1) YFT1 is the oldest shell midden known, providing minimum constraints on full coastal adaptation by ∼120 ka; 2) despite rapid sea-level drop and other climatic changes during occupation, relative shellfish proportions and sizes remain similar, suggesting adaptive foraging along a changing coastline; 3) the YFT1 lithic technocomplex is similar to other west coast assemblages but distinct from potentially synchronous industries along the southern African coast, suggesting human populations were fragmented between seasonal rainfall zones; and 4) accumulation rates (up to 1.8 m/ka) are much higher than previously observed for dated, stratified MSA middens, implying more intense site occupation akin to Later Stone Age middens.

Significance For most of its path through plant bodies, water moves in conduits in the wood. Plant water conduction is crucial for Earth’s biogeochemical cycles, making it important to understand how natural selection shapes conduit diameters along the entire lengths of plant stems. Can mathematical modeling and global sampling explain how wood conduits ought to widen from the tip of a plant to its trunk base? This question is evolutionarily important because xylem conduits should widen in a way that keeps water supply constant to the leaves as a plant grows taller. Moreover, selection should act on economy of construction costs of the conducting system. This issue is ecologically important because it helps suggest why climate change alters vegetation height worldwide., Shaping global water and carbon cycles, plants lift water from roots to leaves through xylem conduits. The importance of xylem water conduction makes it crucial to understand how natural selection deploys conduit diameters within and across plants. Wider conduits transport more water but are likely more vulnerable to conduction-blocking gas embolisms and cost more for a plant to build, a tension necessarily shaping xylem conduit diameters along plant stems. We build on this expectation to present the Widened Pipe Model (WPM) of plant hydraulic evolution, testing it against a global dataset. The WPM predicts that xylem conduits should be narrowest at the stem tips, widening quickly before plateauing toward the stem base. This universal profile emerges from Pareto modeling of a trade-off between just two competing vectors of natural selection: one favoring rapid widening of conduits tip to base, minimizing hydraulic resistance, and another favoring slow widening of conduits, minimizing carbon cost and embolism risk. Our data spanning terrestrial plant orders, life forms, habitats, and sizes conform closely to WPM predictions. The WPM highlights carbon economy as a powerful vector of natural selection shaping plant function. It further implies that factors that cause resistance in plant conductive systems, such as conduit pit membrane resistance, should scale in exact harmony with tip-to-base conduit widening. Furthermore, the WPM implies that alterations in the environments of individual plants should lead to changes in plant height, for example, shedding terminal branches and resprouting at lower height under drier climates, thus achieving narrower and potentially more embolism-resistant conduits.

In nature-based treatment systems, such as constructed wetlands, plant uptake of nutrients can be a significant removal pathway. Current methods for quantifying plant uptake of nitrogen in constructed wetlands, which often involve harvesting biomass and assuming that all nitrogen stored in plants was derived from wastewater, are inappropriate in pilot- and full-scale systems where other sources of nitrogen are available. To improve our understanding of nitrogen cycling in constructed wetlands, we developed a new method to quantify plant uptake of nitrogen by using stable isotopes and a mixing model to distinguish between nitrogen sources. We applied this new method to a pilot-scale horizontal levee system (i.e., a subsurface constructed wetland) over a two-year monitoring period, during which 14% of nitrogen in plants was wastewater-derived on average and the remaining plant nitrogen was obtained from the soil. Analysis of nitrogen isotopes indicated substantial spatial variability in the wetland: 82% of nitrogen in plants within the first 2m of the slope came from wastewater while less than 12% of plant nitrogen in the remainder of the wetland originated from wastewater. By combining these source contributions with remote-sensing derived total biomass measurements, we calculated that 150kgN (95% CI=50kgN, 330kgN) was taken up and retained by plants during the two-year monitoring period, which corresponded to approximately 8% of nitrogen removed in the wetland. Nitrogen uptake followed seasonal trends, increased as plants matured, and varied based on design parameters (e.g., plant types), suggesting that design decisions can impact this removal pathway. This new method can help inform efforts to understand nitrogen cycling and optimize the design of nature-based nutrient control systems.

The environmental fates and consequences of intensive sulfur (S) applications to croplands are largely unknown. In this study, we used S stable isotopes to identify and trace agricultural S from field-to-watershed scales, an initial and timely step toward constraining the modern S cycle. We conducted our research within the Napa River Watershed, California, US, where vineyards receive frequent fungicidal S sprays. We measured soil and surface water sulfate concentrations ([SO4 2−]) and stable isotopes (δ 34S–SO4 2−), which we refer to in combination as the ‘S fingerprint’. We compared samples collected from vineyards and surrounding forests/grasslands, which receive background atmospheric and geologic S sources. Vineyard δ 34S–SO4 2− values were 9.9 ± 5.9‰ (median ± interquartile range), enriched by ∼10‰ relative to forests/grasslands (−0.28 ± 5.7‰). Vineyards also had roughly three-fold higher [SO4 2−] than forests/grasslands (13.6 and 5.0 mg SO4 2−–S l−1, respectively). Napa River δ 34S–SO4 2− values, reflecting the watershed scale, were similar to those from vineyards (10.5 ± 7.0‰), despite vineyard agriculture constituting only ∼11% of the watershed area. Combined, our results provide important evidence that agricultural S is traceable at field-to-watershed scales, a critical step toward determining the consequences of agricultural alterations to the modern S cycle.

In order to apply isotope methods towards enhancing our understanding of water movement through the soil-plant-atmospheric continuum, thye authors first determine what controls water movement through each separate compartment. Water movement in soils and into plant root systems will be a complex function of soil properties, root anatomy and morphology and the stability of water sources which plants may use at different times or from different depths. Root water uptake will also be determined by ‘demand’ by the plant canopy and patterns by which stomata regulate water loss from leaves. The use of stable hydrogen and oxygen isotopes in studies of plant water relations has dramatically improved our ability to link plants to their source water through nondestructive means. Isotopes have also allowed researchers to examine temporal shifts in water use that cannot be determined from documenting the location of roots in the soil profile.

Dew is an important water resource for plants in most deserts. The mechanism that allows desert plants to use dew water was studied using an isotopic water tracer approach. Most plants use water directly from the soil; the roots transfer the water to the rest of the plant, where it is required for all metabolic functions. However, many plants can also take up water into their leaves and stems. Examining the dew water uptake pathways in desert plants can lend insight on another all water-use pathways examination. We determined where and how dew water enters plants in the water limited Negev desert. Highly depleted isotopic water was sprayed on three different dominant plant species of the Negev desert-Artemesia sieberi, Salsola inermis and Haloxylon scoparium-and its entry into the plant was followed. Water was sprayed onto the soil only, or on the leaves/stems only (with soil covered to prevent water entry via root uptake). Thereafter, the isotopic composition of water in the roots and stems were measured at various time points. The results show that each plant species used the dew water to a different extent, and we obtained evidence of foliar uptake capacity of dew water that varied depending on the microenvironmental conditions. A. sieberi took up the greatest amount of dew water through both stems and roots, S. inermis took up dew water mainly from the roots, and H. scoparium showed the least dew capture overall.

Water stored in tree stems (i.e., trunks and branches) is an important contributor to transpiration that can improve photosynthetic carbon gain and reduce the probability of cavitation. However, in tall trees, the capacity to store water may decline with height because of chronically low water potentials associated with the gravitational potential gradient. We quantified the importance of elastic stem water storage in the top 5–6 m of large (4.2–5.0 m diameter at breast height, 82.1–86.3 m tall) Sequoiadendron giganteum (Lindley) J. Buchholz (giant sequoia) trees using a combination of architectural measurements and automated sensors that monitored summertime diel rhythms in sap flow, stem diameter and water potential. Stem water storage contributed 1.5–1.8% of water transpired at the tree tops, and hydraulic capacitance ranged from 2.6 to 4.1 l MPa−1 m−3. These values, which are considerably smaller than reported for shorter trees, may be associated with persistently low water potentials imposed by gravity and could indicate a trend of decreasing water storage dynamics with height in tree. Branch diameter contraction and expansion consistently and substantially lagged behind fluxes in water potential and sap flow, which occurred in sync. This lag suggests that the inner bark, which consists mostly of live secondary phloem tissue, was an important hydraulic capacitor, and that hydraulic resistance between xylem and phloem retards water transfer between these tissues. We also measured tree-base sap flux, which lagged behind that measured in trunks near the tree tops, indicating additional storage in the large trunks between these measurement positions. Whole-tree sap flow ranged from 2227 to 3752 l day−1, corroborating previous records for similar-sized giant sequoia and representing the largest yet reported for any individual tree. Despite such extraordinarily high daily water use, we estimate that water stored in tree-top stems contributes minimally to transpiration on typical summer days.

Climate warming is facilitating the expansion of many cold-sensitive woody species in woodland-grassland ecotones worldwide. Recent research has demonstrated that this range expansion can be further enhanced by positive vegetation-microclimate feedbacks whereby woody canopies induce local nocturnal warming, which reduces freeze-induced damage and favors the establishment of woody plants. However, this local positive feedback can be counteracted by biotic drivers such as browsing and the associated consumption of shrub biomass. The joint effects of large-scale climate warming and local-scale microclimate feedbacks on woody vegetation dynamics in these ecotones remain poorly understood. Here, we used a combination of experimental and modeling approaches to investigate the effects of woody cover on microclimate and the consequent implications on ecological stability in North American coastal ecosystems. We found greater browsing pressure and significant warming (~2°C) beneath shrub canopies compared to adjacent grasslands, which reduces shrub seedlings' exposure to cold damage. Cold sensitivity is evidenced by the significant decline in xylem hydraulic conductivity in shrub seedlings when temperatures dropped below -2°C. Despite the negative browsing-vegetation feedback, a small increase in minimum temperature can induce critical transitions from grass to woody plant dominance. Our framework also predicts the threshold temperature of -7°C for mangrove-salt marsh ecotones on the Atlantic coast of Florida. Above this reference temperature a critical transition may occur from salt marsh to mangrove vegetation, in agreement with empirical studies. Thus, the interaction between ongoing global warming trends and microclimate feedbacks may significantly alter woody vegetation dynamics and ecological stability in coastal ecosystems where woody plant expansion is primarily constrained by extreme low temperature events.

The large-scale patterns of movement for the Sharp-shinned Hawk (Accipiter striatus), a small forest hawk found throughout western North America, are largely unknown. However, based on field observations we set out to test the hypothesis that juvenile migratory A. striatus caught along two distinct migration routes on opposite sides of the Sierra Nevada Mountains of North America (Pacific Coast and Intermountain Migratory Flyways) come from geographically different natal populations. We applied stable isotope analysis of hydrogen (H) and oxygen (O) of feathers, and large scale models of spatial isotopic variation (isoscapes) to formulate spatially explicit predictions of the origin of the migrant birds. Novel relationships were assessed between the measured hydrogen and oxygen isotope values of feathers from A. striatus museum specimens of known origin and the isoscape modeled hydrogen and oxygen isotope values of precipitation at those known locations. We used these relationships to predict the origin regions for birds migrating along the two flyways from the measured isotope values of migrant’s feathers and the associated hydrogen and oxygen isotopic composition of precipitation where these feathers were formed. The birds from the two migration routes had overlap in their natal/breeding origins and did not differentiate into fully separate migratory populations, with birds from the Pacific Coast Migratory Flyway showing broader natal geographic origins than those from the Intermountain Flyway. The methodology based on oxygen isotopes had, in general, less predictive power than the one based on hydrogen. There was broad agreement between the two isotope approaches in the geographic assignment of the origins of birds migrating along the Pacific Coast Flyway, but not for those migrating along the Intermountain Migratory Flyway. These results are discussed in terms of their implications for conservation efforts of A. striatus in western North America, and the use of combined hydrogen and oxygen stable isotope analysis to track the movement of birds of prey on continental scales.

© 2018 The US and Mexico share a common history in many areas, including language and culture. They face ecological changes due to the increased frequency and severity of droughts and rising energy demands; trends that entail economic costs for both nations and major implications for human wellbeing. We describe an ongoing effort by the Environment Working Group (EWG), created by The University of California's UC-Mexico initiative in 2015, to promote binational research, teaching, and outreach collaborations on the implications of climate change for Mexico and California. We synthesize current knowledge about the most pressing issues related to climate change in the US-Mexico border region and provide examples of cross-border discoveries and research initiatives, highlighting the need to move forward in six broad rubrics. This and similar binational cooperation efforts can lead to improved living standards, generate a collaborative mindset among participating universities, and create an international network to address urgent sustainability challenges affecting both countries.

To explore the role of past environmental changes in shaping interspecific differences in genetic structure among tuco-tucos (Rodentia: Ctenomyidae, Ctenomys) from the Limay Valley of northern Patagonia, we compared δ13C and δ18O values for modern and historical tooth enamel from the colonial tuco-tuco (C. sociabilis) and the parapatric Patagonian tuco-tuco (C. haigi). Specifically, we sought to test the hypothesis that a mid-Holocene loss of genetic variation in C. sociabilis was associated with environmental change. For both species, the δ13C composition of tooth enamel differed significantly (~ 5.1? difference in mean δ13C values) between modern and Holocene (0.5?12 kyr B.P.) samples, suggesting marked temporal changes in the vegetation consumed. For C. sociabilis, no statistically significant differences in the δ13C composition of teeth were found among individuals from the early, middle, and late Holocene. In contrast, significant differences in δ13C values were found between middle and late Holocene samples from C. haigi. None of our analyses revealed statistically significant differences in δ18O values. Thus, although the diets of both study species have changed from the Holocene to the present, only C. haigi experienced significant dietary changes during the Holocene. We suggest that the ecologically more specialized C. sociabilis continued to consume the same food resources even as changing Holocene conditions caused preferred habitats to disappear. In contrast, the more generalized C. haigi altered its diet, allowing populations of this species to remain widespread despite changing Holocene environments. Fil: Takenaka, Risa. University of California at Berkeley; Estados Unidos Fil: Miller, Melanie. University of Otago; Nueva Zelanda. University of California at Berkeley; Estados Unidos Fil: Tammone, Mauro Nicolás. Administración de Parques Nacionales. Parque Nacional Nahuel Huapi. Programa de Estudios Aplicados a la Conservación; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: Dawson, Todd. University of California at Berkeley; Estados Unidos Fil: Lacey, Eileen Anne. University of California at Berkeley; Estados Unidos

The large-scale patterns of movement for the Sharp-shinned Hawk (Accipiter striatus), a small forest hawk found throughout western North America, are largely unknown. However, based on field observations we set out to test the hypothesis that juvenile migratory A. striatus caught along two distinct migration routes on opposite sides of the Sierra Nevada Mountains of North America (Pacific Coast and Intermountain Migratory Flyways) come from geographically different natal populations. We applied stable isotope analysis of hydrogen (H) and oxygen (O) of feathers, and large scale models of spatial isotopic variation (isoscapes) to formulate spatially explicit predictions of the origin of the migrant birds. Novel relationships were assessed between the measured hydrogen and oxygen isotope values of feathers from A. striatus museum specimens of known origin and the isoscape modeled hydrogen and oxygen isotope values of precipitation at those known locations. We used these relationships to predict the origin regions for birds migrating along the two flyways from the measured isotope values of migrant’s feathers and the associated hydrogen and oxygen isotopic composition of precipitation where these feathers were formed. The birds from the two migration routes had overlap in their natal/breeding origins and did not differentiate into fully separate migratory populations, with birds from the Pacific Coast Migratory Flyway showing broader natal geographic origins then those from the Intermountain Flyway. The methodology based on oxygen isotopes had, in general, less predictive power than the one based on hydrogen. There was broad agreement between the two isotope approaches in the geographic assignment of the origins of birds migrating along the Pacific Coast Flyway, but not for those migrating along the Intermountain Migratory Flyway. These results are discussed in terms of their implications for conservation efforts of A. striatus in western North America, and the use of combined hydrogen and oxygen stable isotope analysis to track the movement of birds of prey on continental scales.

The emergence of critical zone (CZ) science has provided an integrative platform for investigating plant ecophysiology in the context of landscape evolution, weathering and hydrology. The CZ lies between the top of the vegetation canopy and fresh, chemically unaltered bedrock and plays a pivotal role in sustaining life. We consider what the CZ perspective has recently brought to the study of plant ecophysiology. We specifically highlight novel research demonstrating the importance of the deeper subsurface for plant water and nutrient relations. We also point to knowledge gaps and research opportunities, emphasising, in particular, greater focus on the roles of deep, nonsoil resources and how those resources influence and coevolve with plants as a frontier of plant ecophysiological research.

California experienced severe drought from 2012 to 2016, and there were visible changes in the forest canopy throughout the State. In 2014, unprecedented foliage dieback was recorded in giant sequoia (Sequoiadendron giganteum) trees in Sequoia National Park, in the southern California Sierra Nevada mountains. Although visible changes in sequoia canopies can be recorded, biochemical and physiological responses to drought stress in giant sequoia canopies are not well understood. Ground-based measurements provide insight into the mechanisms of drought responses in trees, but are often limited to few individuals, especially in trees of tall stature such as giant sequoia. Recent studies demonstrate that remotely measured forest canopy water content (CWC) is a general indicator of canopy response to drought, but the underpinning leaf- to canopy-level causes of observed variation in CWC remain poorly understood. We combined field and airborne remote sensing measurements taken in 2015 and 2016 to assess the biophysical responses of giant sequoias to drought. In 49 study trees, CWC was related to leaf water potential, but not to the other foliar traits, suggesting that changes in CWC were made at whole-canopy rather than leaf scales. We found a non-random, spatially varying pattern in mapped CWC, with lower CWC values at lower elevation and along the outer edges of the groves. This pattern was also observed in empirical measurements of foliage dieback from the ground, and in mapped CWC across multiple sequoia groves in this region, supporting the hypothesis that drought stress is expressed in canopy-level changes in giant sequoias. The fact that we can clearly detect a relationship between CWC and foliage dieback, even without taking into account prior variability or new leaf growth, strongly suggests that remotely sensed CWC, and changes in CWC, are a useful measure of water stress in giant sequoia, and valuable for assessing and managing these iconic forests in drought.

Hotter droughts are becoming more common as climate change progresses, and they may already have caused instances of forest dieback on all forested continents. Learning from hotter droughts, including where on the landscape forests are more or less vulnerable to these events, is critical to help resource managers proactively prepare for the future. As part of our Leaf to Landscape Project, we measured the response of giant sequoia, the world’s largest tree species, to the extreme 2012–2016 hotter drought in California. The project integrated leaf-level physiology measurements, crown-level foliage dieback surveys, and remotely sensed canopy water content (CWC) to shed light on mechanisms and spatial patterns in drought response. Here we summarize initial findings, present a conceptual model of drought response, and discuss management implications; details are presented in the other four articles of the special section on Giant Sequoias and Drought. Giant sequoias exhibited both leaf- and canopy-level responses that were effective in protecting whole-tree hydraulic integrity for the vast majority of individual sequoias. Very few giant sequoias died during the drought compared to other mixed conifer tree species; however, the magnitude of sequoia drought response varied across the landscape. This variability was partially explained by local site characteristics, including variables related to site water balance. We found that low CWC is an indicator of recent foliage dieback, which occurs when stress levels are high enough that leaf-level adjustments alone are insufficient for giant sequoias to maintain hydraulic integrity. CWC or change in CWC may be useful indicators of drought stress that reveal patterns of vulnerability to future hotter droughts. Future work will measure recovery from the drought and strengthen our ability to interpret CWC maps. Our ultimate goal is to produce giant sequoia vulnerability maps to help target management actions, such as reducing other stressors, increasing resistance to hotter drought through prescribed fire or mechanical thinning, and planting sequoias in projected future suitable habitat, which may occur outside current grove distributions. We suggest that managers compare different types of vulnerability assessments and combine vulnerability maps with other sources of information to inform decisions.

Drought is expected to become an increasingly important stressor on forests globally, and understanding the physiological mechanisms driving tree drought response is essential for developing effective mitigation and conservation measures for these ecosystems. In 2014, during California’s 2012–2016 “hotter” drought in which higher temperatures exacerbated the effects of low water availability, many giant sequoia trees in the Sierra Nevada mountains exhibited foliage dieback at levels previously unreported. We hypothesized that this apparent drought-induced foliage dieback was associated with spatial patterns of site water balance and consequently tree water status and physiology. As part of an ongoing collaborative project aimed at understanding and mapping giant sequoia drought response and vulnerability at leaf to landscape scales, in 2015, 2016 and 2017 we climbed and measured leaf-level water status and physiology of mature and seedling giant sequoia trees at sites exhibiting low and high average foliage dieback in Giant Forest, Sequoia National Park. We also compared our 2015–2017 measurements with similar measurements made in giant sequoias prior to the commencement of the 2012–2016 drought. We found that during the drought, leaf water potentials of both mature and seedling sequoias were as low or lower than any previous measurements, and leaf water potentials of some seedlings were as low as the tops of mature trees. In contrast to our expectations, we found similar water potentials in both foliage dieback classes, although there was a high degree of within-site variability and in some measurement periods trees and seedlings growing in flat or meadow-side topographic positions had higher water potentials than those growing on mid- or upslope positions with presumably less favorable water supply. Leaf-level adjustments included multiple mechanisms to reduce water loss and resist the effects of desiccation at both seasonal and annual time scales, including stomatal closure, redistribution of leaf water to less mobile storage, and a shift in leaf carbon fractions to build tougher, more drought-resistant foliage. Our results suggest that giant sequoia employs a drought avoidance water regulation strategy and leaf-level adjustments are sufficient to maintain relatively isohydric water potentials above critical thresholds under most conditions. Because of the high severity of the 2012–2016 drought, however, the capacity for leaf-level compensation in some giant sequoias was overwhelmed and both leaf- and crown-level adjustments were necessary to maintain favorable water status to protect whole-tree hydraulic functioning. Additional research aimed at understanding and monitoring the consequences of future hotter droughts is necessary in order to determine the long-term persistence of giant sequoias within their current geographic distribution.

Contents Summary 1156 I. Introduction 1156 II. How often are leaves wet? 1157 III. The costs of leaf wetting 1157 IV. The real and potential benefits of leaf wetting 1161 V. Wet leaves: costs, benefits and tradeoffs in a changing world 1165 Acknowledgements 1166 References 1166 SUMMARY: An often-overlooked feature of all plants is that their leaf surfaces are wet for significant periods over their lifetimes. Leaf wetting has a number of direct and indirect effects on plant function from the scale of the leaf to that of the ecosystem. The costs of leaf wetting for plant function, such as the growth of pathogens and the leaching of nutrients, have long been recognized. However, an emerging body of research has also begun to demonstrate some very clear benefits. For instance, leaf wetting can improve plant-water relations and lead to increased photosynthesis. Leaf wetting may also lead to synergistic effects on plant function, such as when leaf water potential improvements lead to enhanced growth that does not occur when plant leaves are dry. We identify important reasons why leaf wetting can be critical for plant sciences to not only acknowledge, but also directly address, in future research. To do so, we provide a framework for the consideration of the relative balance of the various costs and benefits resulting from leaf wetting, as well as how this balance may be expected to change given projected scenarios of global climate change in the future.

Environmental changes have resulted in significant declines in native riparian forests that are comprised largely of dioecious tree taxa, including boxelder and iconic cottonwood/willow gallery forests. Dioecious species may be especially vulnerable to the effects of climate change given that they often exhibit skewed sex ratios that are reinforced by physiological and morphological specialization of each sex to specific microhabitats. A comprehensive data synthesis suggests that male individuals of boxelder and cottonwood taxa have a higher representation on dry microhabitats than females and are less physiologically sensitive to increased aridity than co-occurring females. Consequently, extreme male-biased sex ratios are possible under future climate conditions that could reduce population fitness below a sustainable threshold. Riparian willows, on the other hand, generally do not express obvious sexual dimorphism in habitat preference or physiological sensitivity to aridity. Thus, it is unclear whether climate change will impact population structure of willows in ways that parallel other dioecious riparian tree taxa. Future riparian tree restoration programs should aim to maintain future sex ratio balance that maximizes population fitness under projected hydro-climatological conditions. Recent advances in genomics will likely provide the critical tools for early sex determination in pre-reproductive trees across riparian tree species such that sex ratio balance could be targeted during initial stages of restoration, along with adaptations for drought tolerance and other key traits that are essential for survival under future conditions.

To understand drivers of hydroclimate variability in north-coastal California, we obtained tree cross-sections from eleven coastal redwoods (mean age of 1232 years old) from the northern half of the species range. Tree rings from eight trees were cross-dated and sampled at sub-annual resolution for carbon isotope discrimination (Δ13C) and oxygen isotope composition (δ18O). Tree-ring Δ13C and δ18O, compared to modern climate data, demonstrate these signals primarily record summertime hydroclimate variability—primarily through variables associated with evaporative conditions and/or precipitation. Our 1100-year stable isotope chronologies showed that north-coastal California did not undergo the megadroughts observed elsewhere in California and the western United States. This result implicates extended periods of low winter precipitation, rather than growing season evaporation, as the primary driver of previous megadroughts across California and neighboring regions. Compared to cool conditions prevailing over the Northern Hemisphere during the Little Ice age (1301–1875 of the common era, CE), the frequency of isotopic events of a certain magnitude was greater during periods with warmer Northern Hemisphere temperatures such as the Medieval Climate Anomaly (900–1300 CE) and the modern period (1876 to present). This association between tree-ring isotopic variability and long-term shifts in temperatures is consistent with the expected patterns in mid-latitude hydroclimate variability expected from arctic amplification (i.e., shifts in equator-to-pole temperature differences that modify jet stream speed and amplitude) or amplified quasi-resonant wave activity (i.e., wave-patterns in high-altitude winds that become “trapped” within a certain pattern, thereby producing a longer-duration periods of drought or wetness) across mid-latitudes during the boreal summer.