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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.

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.

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.

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 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.

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.

Author(s): Campbell, JE; Whelan, ME; Berry, JA; Hilton, TW; Zumkehr, A; Stinecipher, J; Lu, Y; Kornfeld, A; Seibt, U; Dawson, TE; Montzka, SA; Baker, IT; Kulkarni, S; Wang, Y; Herndon, SC; Zahniser, MS; Commane, R; Loik, ME | Abstract: The future resilience of coast redwoods (Sequoia sempervirens) is now of critical concern due to the detection of a 33% decline in California coastal fog over the 20th century. However, ecosystem-scale measurements of photosynthesis and stomatal conductance are challenging in coast redwood forests, making it difficult to anticipate the impacts of future changes in fog. To address this methodological problem, we explore coastal variations in atmospheric carbonyl sulfide (COS or OCS), which could potentially be used as a tracer of these ecosystem processes. We conducted atmospheric flask campaigns in coast redwood sites, sampling at surface heights and in the canopy (~70nm), at the University of California Landels-Hill Big Creek Reserve and Big Basin State Park. We simulated COS atmosphere-biosphere exchange with a high-resolution 3-D model to interpret these data. Flask measurements indicated a persistent daytime drawdown between the coast and the downwind forest (45n±n6nppt COS) that is consistent with the expected relationship between COS plant uptake, stomatal conductance, and gross primary production. Other sources and sinks of COS that could introduce noise to the COS tracer technique (soils, anthropogenic activity, nocturnal plant uptake, and surface hydrolysis on leaves) are likely to be small relative to daytime COS plant uptake. These results suggest that COS measurements may be useful for making ecosystem-scale estimates of carbon, water, and energy exchange in coast redwood forests.

© 2017 John Wiley & Sons Ltd Climate, physical landscapes, and biota interact to generate heterogeneous hydrologic conditions in space and over time, which are reflected in spatial patterns of species distributions. As these species distributions respond to rapid climate change, microrefugia may support local species persistence in the face of deteriorating climatic suitability. Recent focus on temperature as a determinant of microrefugia insufficiently accounts for the importance of hydrologic processes and changing water availability with changing climate. Where water scarcity is a major limitation now or under future climates, hydrologic microrefugia are likely to prove essential for species persistence, particularly for sessile species and plants. Zones of high relative water availability – mesic microenvironments – are generated by a wide array of hydrologic processes, and may be loosely coupled to climatic processes and therefore buffered from climate change. Here, we review the mechanisms that generate mesic microenvironments and their likely robustness in the face of climate change. We argue that mesic microenvironments will act as species-specific refugia only if the nature and space/time variability in water availability are compatible with the ecological requirements of a target species. We illustrate this argument with case studies drawn from California oak woodland ecosystems. We posit that identification of hydrologic refugia could form a cornerstone of climate-cognizant conservation strategies, but that this would require improved understanding of climate change effects on key hydrologic processes, including frequently cryptic processes such as groundwater flow.

Drought extent and severity have increased and are predicted to continue to increase in many parts of the world. Understanding tree vulnerability to drought at both individual and species levels is key to ongoing forest management and preparation for future transitions in community composition. The influence of subsurface hydrologic processes is particularly important in water-limited ecosystems, and is an under-studied aspect of tree drought vulnerability. With California's 2013-2016 extraordinary drought as a natural experiment, we studied four co-occurring woodland tree species, blue oak (Quercus douglasii), valley oak (Quercus lobata), gray pine (Pinus sabiniana), and California juniper (Juniperus californica), examining drought vulnerability as a function of climate, lithology and hydrology using regional aerial dieback surveys and site-scale field surveys. We found that in addition to climatic drought severity (i.e., rainfall), subsurface processes explained variation in drought vulnerability within and across species at both scales. Regionally for blue oak, severity of dieback was related to the bedrock lithology, with higher mortality on igneous and metamorphic substrates, and to regional reductions in groundwater. At the site scale, access to deep subsurface water, evidenced by stem water stable isotope composition, was related to canopy condition across all species. Along hillslope gradients, channel locations supported similar environments in terms of water stress across a wide climatic gradient, indicating that subsurface hydrology mediates species' experience of drought, and that areas associated with persistent access to subsurface hydrologic resources may provide important refugia at species' xeric range edges. Despite this persistent overall influence of the subsurface environment, individual species showed markedly different response patterns. We argue that hydrologic niche segregation can be a useful lens through which to interpret these differences in vulnerability to climatic drought and climate change.

Many animals are considered to be specialists because they have feeding structures that are fine-tuned for consuming specific prey. For example, "smasher" mantis shrimp have highly specialized predatory appendages that generate forceful strikes to break apart hard-shelled prey. Anecdotal observations suggest, however, that the diet of smashers may include soft-bodied prey as well. Our goal was to examine the diet breadth of the smasher mantis shrimp, Neogonodactylus bredini, to determine whether it has a narrow diet of hard-shelled prey. We combined studies of prey abundance, feeding behavior, and stable isotope analyses of diet in both seagrass and coral rubble to determine if N. bredini's diet was consistent across different habitat types. The abundances of hard-shelled and soft-bodied prey varied between habitats. In feeding experiments, N. bredini consumed both prey types. N. bredini consumed a range of different prey in the field as well and, unexpectedly, the stable isotope analysis demonstrated that soft-bodied prey comprised a large proportion (29-53 %) of the diet in both habitats. Using a Bayesian mixing model framework (MixSIAR), we found that this result held even when we used uninformative, or generalist, priors and informative priors reflecting a specialist diet on hard-shelled prey and prey abundances in the field. Thus, contrary to expectation, the specialized feeding morphology of N. bredini corresponds to a broad diet of both hard-shelled and soft-bodied prey. Using multiple lines of study to describe the natural diets of other presumed specialists may demonstrate that specialized morphology often broadens rather than narrows diet breadth.

In this commentary, we build on discussions that emerged during the workshop "Isotope-based studies of water partitioning and plant-soil interactions in forested and agricultural environments" held in San Casciano Val di Pesa, Italy, in September 2017. Quantifying and understanding how water cycles through the Earth's critical zone is important to provide society and policy makers with the scientific background to manage water resources sustainably, especially considering the ever-increasing worldwide concern about water scarcity. Stable isotopes of hydrogen and oxygen in water have proven to be a powerful tool to track water fluxes in the critical zone. However, both mechanistic complexities (e.g., mixing and fractionation processes, heterogeneity of natural systems) and practical methodological issues (e.g., lack of standard protocols to sample specific compartments, such as soil water and xylem water) limit the application of stable water isotopes in critical zone science. In this commentary, we examine some of the opportunities and critical challenges of using isotope-based ecohydrological applications, and outline new perspectives focused on interdisciplinary research opportunities for this important tool in water and environmental science.

Cross-boundary transfers of nutrients can profoundly shape the ecology of recipient systems. The common hippopotamus, Hippopotamus amphibius, is a significant vector of such subsidies from terrestrial to river ecosystems. We compared river pools with high and low densities of H. amphibius to determine how H. amphibius subsidies shape the chemistry and ecology of aquatic communities. Our study watershed, like many in sub-Saharan Africa, has been severely impacted by anthropogenic water abstraction reducing dry-season flow to zero. We conducted observations for multiple years over wet and dry seasons to identify how hydrological variability influences the impacts of H. amphibius. During the wet season, when the river was flowing, we detected no differences in water chemistry and nutrient parameters between pools with high and low densities of H. amphibius. Likewise, the diversity and abundance of fish and aquatic insect communities were indistinguishable. During the dry season, however, high-density H. amphibius pools differed drastically in almost all measured attributes of water chemistry and exhibited depressed fish and insect diversity and fish abundance compared with low-density H. amphibius pools. Scaled up to the entire watershed, we estimate that H. amphibius in this hydrologically altered watershed reduces dry-season fish abundance and indices of gamma-level diversity by 41% and 16%, respectively, but appears to promote aquatic invertebrate diversity. Widespread human-driven shifts in hydrology appear to redefine the role of H. amphibius, altering their influence on ecosystem diversity and functioning in a fashion that may be more severe than presently appreciated.

The Hawaiian Islands offer a unique opportunity to test how changes in the properties of an isolated ecosystem are propagated through the organisms that occur within that ecosystem. The age-structured arrangement of volcanic-derived substrates follows a regular progression over space and, by inference, time. We test how well documented successional changes in soil chemistry and associated vegetation are reflected in organisms at higher trophic levels—specifically, predatory arthropods (spiders)—across a range of functional groups. We focus on three separate spider lineages: one that builds capture webs, one that hunts actively, and one that specializes on eating other spiders. We analyze spiders from three sites across the Hawaiian chronosequence with substrate ages ranging from 200 to 20,000 years. To measure the extent to which chemical signatures of terrestrial substrates are propagated through higher trophic levels, we use standard stable isotope analyses of nitrogen and carbon, with plant leaves included as a baseline. The target taxa show the expected shift in isotope ratios ofδ15N with trophic level, from plants to cursorial spiders to web-builders to spider eaters. Remarkably, organisms at all trophic levels also precisely reflect the successional changes in the soil stoichiometry of the island chronosequence, demonstrating how the biogeochemistry of the entire food web is determined by ecosystem succession of the substrates on which the organisms have evolved.

Author(s): Templer, PH; Weathers, KC; Ewing, HA; Dawson, TE; Mambelli, S; Lindsey, AM; Webb, J; Boukili, VK; Firestone, MK | Abstract: A defining feature of the redwood forest in coastal California is the presence of fog in the summer months, a time when there is typically little rainfall. Our goal was to determine the role of summer fog in canopy transformation of nitrogen, nitrogen uptake by trees and photosynthesis within a coastal redwood forest ecosystem. We measured horizontal and vertical inputs of nitrogen, the isotopic composition of nitrogen in a variety of atmospheric sources (summer fog, winter rain and throughfall throughout the year), nitrogen pools (soil solution) and plant tissue (roots and foliage), as well as rates of photosynthesis and nitrogen uptake by trees. Throughfall nitrogen fluxes were greater at the forest edge compared to the interior both within the canopy (sampled 10m above-ground) and onto the forest floor (sampled 1m above-ground; Pl0.05). Similarly, soil solution NO3- and total inorganic nitrogen were greater at the forest edge compared to the interior (P=0.0014 and 0.009, respectively). Whereas natural abundance δ15NO3 values were not significantly different between winter rain (measured as bulk precipitation) and summer fog water (average δ15N=-1.2±0.680/00), δ15NH4 values were significantly greater in fog water (11.4±2.70/00) compared to rain (1.2±0.90/00). We found no difference in δ15N in roots from forest edge trees compared to interior trees. In contrast, nitrogen concentrations and δ15N in foliage from forest edge trees were significantly greater compared to interior trees (Pl0.0001), suggesting that the leaves of forest edge trees may be obtaining a greater proportion of their nitrogen from fog compared to those of the interior trees. Natural abundance 13C of leaf sugars and rates of photosynthesis were significantly higher at the forest edge compared to the interior during the fog season (Pl0.05), but not different between locations in the rain season (Pg0.05). Nitrification in the forest floor, rather than the canopy, is the primary source of NO3- in these soils throughout the year. Synthesis. Summer fog provides nitrogen directly and indirectly to redwood trees, especially those at the forest edge, and affects the physiologic function of redwood trees. Summer fog provides nitrogen directly and indirectly to redwood trees, especially those at the forest edge, and affects the physiologic function of redwood trees.

The ecological importance of the common hippopotamus (Hippopotamus amphibius) in aquatic ecosystems is becoming increasingly well known. These unique megaherbivores are also likely to have a formative influence on the terrestrial ecosystems in which they forage. In this study, we employed a novel exclosure design to exclude H. amphibius from experimental plots on near-river grasslands. Our three-year implementation of this experiment revealed a substantial influence of H. amphibius removal on both plant communities and soil chemistry. H. amphibius significantly reduced grassland canopy height, increased the leafiness of common grasses, reduced woody plant abundance and size, and increased the concentrations of several soil elements. Many of the soil chemistry changes that we experimentally induced by exclusion of H. amphibius were mirrored in the soil chemistry differences between naturally occurring habitats of frequent (grazing lawns) and infrequent (shrub forest) use by H. amphibius and other grazing herbivores. In contrast to existing hypotheses regarding grazing species, we found that H. amphibius had little effect on local plant species richness. Simultaneous observations of exclosures designed to remove all large herbivores revealed that H. amphibius removal had ecologically significant impacts, but that the removal of all species of large herbivores generated more pronounced impacts than the removal of H. amphibius alone. In aggregate, our results suggest that H. amphibius have myriad effects on their terrestrial habitats that likely improve the quality of forage available for other herbivores. We suggest that ongoing losses of this vulnerable megaherbivore are likely to cause significant ecological change.

It is commonly assumed that transpiration does not occur at night because leaf stomata are closed in the dark. We tested this assumption across a diversity of ecosystems and woody plant species by various methods to explore the circumstances when this assumption is false. Our primary goals were: (1) to evaluate the nature and magnitude of nighttime transpiration, E(n), or stomatal conductance, g(n); and (2) to seek potential generalizations about where and when it occurs. Sap-flow, porometry and stable isotope tracer measurements were made on 18 tree and eight shrub species from seven ecosystem types. Coupled with environmental data, our findings revealed that most of these species transpired at night. For some species and circumstances, nighttime leaf water loss constituted a significant fraction of total daily water use. Our evidence shows that E(n) or g(n) can occur in all but one shrub species across the systems we investigated. However, under conditions of high nighttime evaporative demand or low soil water availability, stomata were closed and E(n) or g(n) approached zero in eleven tree and seven shrub species. When soil water was available, E(n) or g(n) was measurable in these same species demonstrating plasticity for E(n) or g(n). We detected E(n) or g(n) in both trees and shrubs, and values were highest in plants from sites with higher soil water contents and in plants from ecosystems that were less prone to atmospheric or soil water deficits. Irrespective of plant or ecosystem type, many species showed E(n) or g(n) when soil water deficits were slight or non-existent, or immediately after rainfall events that followed a period of soil water deficit. The strongest relationship was between E(n) or g(n) and warm, low humidity and (or) windy (> 0.8 m s(-1)) nights when the vapor pressure deficit remained high (> 0.2 kPa in wet sites, > 0.7 kPa in dry sites). Why E(n) or g(n) occurs likely varies with species and ecosystem type; however, our data support four plausible explanations: (1) it may facilitate carbon fixation earlier in the day because stomata are already open; (2) it may enhance nutrient supply to distal parts of the crown when these nutrients are most available (in wet soils) and transport is rapid; (3) it may allow for the delivery of dissolved O(2) via the parenchyma to woody tissue sinks; or (4) it may occur simply because of leaky cuticles in older leaves or when stomata cannot close fully because of obstructions from stomatal (waxy) plugs, leaf endophytes or asymmetrical guard cells (all non-adaptive reasons). We discuss the methodological, ecophysiological, and theoretical implications of the occurrence of E(n) or g(n) for investigations at a variety of scales.

Hydrogen-isotopic abundances of lipid biomarkers are emerging as important proxies in the study of ancient environments and ecosystems. A decade ago, pioneering studies made use of new analytical methods and demonstrated that the hydrogen-isotopic composition of individual lipids from aquatic and terrestrial organisms can be related to the composition of their growth (i.e., environmental) water. Subsequently, compound-specific deuterium/hydrogen (D/H) ratios of sedimentary biomarkers have been increasingly used as paleohydrological proxies over a range of geological timescales. Isotopic fractionation observed between hydrogen in environmental water and hydrogen in lipids, however, is sensitive to biochemical, physiological, and environmental influences on the composition of hydrogen available for biosynthesis in cells. Here we review the factors and processes that are known to influence the hydrogen-isotopic compositions of lipids—especially n-alkanes—from photosynthesizing organisms, and we provide a framework for interpreting their D/H ratios from ancient sediments and identify future research opportunities.

Coast redwood (Sequoia sempervirens) ecosystems are strongly influenced by the presence of summer marine fog, and variation in fog frequency is closely linked to climate variation in the NE Pacific region. Because oxygen isotope composition (δ18O) of organic matter records distinct water sources (e.g. summertime fog vs. winter precipitation) and carbon isotopes (δ13C) are typically sensitive to humidity and water status, it then follows that inter-annual variation in tree-ring isotope ratios, which are coherent across multiple sites, should preserve a potentially powerful proxy for climate reconstruction. Here we present an analysis of a 50-year time series for both δ18O and δ13C values from subdivided tree rings obtained from multiple redwood trees at multiple sites. Within-site and between-site correlations were highly significant (p < 0.01) for the δ18O time series indicating a regionally coherent common forcing of δ18O fractionation. Within-site and between-site correlation coefficients were ...

The stable hydrogen isotope ratios (δD) of leaf wax n-alkanes record valuable information on plant and ecosystem water relations. It remains, however, unknown if leaf wax n-alkane δD values record only environmental variation during the brief period of time of leaf growth or if leaf wax n-alkane δD values are affected by environmental variability throughout the entire lifespan of a leaf. To resolve these uncertainties, we irrigated Populus trichocarpa trees with a pulse of deuterium-enriched water and used compound-specific stable hydrogen isotope analyses to test if the applied tracer could be recovered from leaf wax n-alkanes of leaves that were at different stages of their development during the tracer application. Our experiment revealed that only leaf wax n-alkanes from leaves that had developed during the time of the tracer application were affected, while leaves that were already fully matured at the time of the tracer application were not. We conclude from our study that under controlled environmental conditions, leaf wax n-alkanes are synthesized only early in the ontogeny of a leaf. Our experiment has implications for the interpretation of leaf wax n-alkane δD values in an environmental context, as it suggests that these compounds record only a brief period of the environmental variability that a leaf experiences throughout its life.

Nitrogen availability and N-cycling dynamics across ecosystems play a critical role in plant functioning and species distribution. Measurements of 15N natural abundance provides a way to assess ecosystem N dynamics, and the range of nitrogen stable isotope values (δ15N) for plants in an ecosystem can indicate divergent strategies for N uptake. We tested the hypotheses that the N-rich seasonally dry forest would have higher soil and leaf δ15N and a smaller range of leaf δ15N values compared to the N-poor cerradao (savanna woodland). We measured N concentration and δ15N in two soil depths and leaves of 27 woody species in cerradao and 26 in seasonally dry forest. As expected, total soil N concentration decreased while soil δ15N value increased with soil depth. Regardless of soil depth, seasonally dry forest soils had higher δ15N and total N concentration compared to cerradao soils. Foliar δ15N values varied from −6.4‰ to 5.9‰ in cerradao and from −2.3‰ to 8.4‰ in seasonally dry forest plants. Phylogenetically independent contrasts analysis and comparisons of δ15N mean values of the most abundant species and species co-occurring in both sites confirmed the hypothesis of higher δ15N for seasonally dry forest in comparison to cerradao. These results corroborate the expectation of higher soil and leaf δ15N values in sites with higher soil N availability. However, except for the most abundant species, no across-site leaf–soil (δ15N leaf –δ15N soil) differences (Δδ15N) were found suggesting that differences in leaf δ15N between cerradao and seasonally dry forest are driven by differences in soil δ15N. Variation of leaf δ15N was large in both sites and only slightly higher in cerradao, suggesting high diversity of N use strategies for both cerradao and seasonally dry forest communities.

Cellulose in plants contains oxygen that derives in most cases from precipitation. Because the stable oxygen isotope composition, δ 18 O, of precipitation is associated with environmental conditions, cellulose δ 18 O should be as well. However, plant physiological models using δ 18 O suggest that cellulose δ 18 O is influenced by a complex mix of both climatic and physiological drivers. This influence complicates the interpretation of cellulose δ 18 O values in a paleo-context. Here, we combined empirical data analyses with mechanistic model simulations to i ) quantify the impacts that the primary climatic drivers humidity ( e a ) and air temperature ( T air ) have on cellulose δ 18 O values in different tropical ecosystems and ii ) determine which environmental signal is dominating cellulose δ 18 O values. Our results revealed that e a and T air equally influence cellulose δ 18 O values and that distinguishing which of these factors dominates the δ 18 O values of cellulose cannot be accomplished in the absence of additional environmental information. However, the individual impacts of e a and T air on the δ 18 O values of cellulose can be integrated into a single index of plant-experienced atmospheric vapor demand: the leaf-to-air vapor pressure difference (VPD). We found a robust relationship between VPD and cellulose δ 18 O values in both empirical and modeled data in all ecosystems that we investigated. Our analysis revealed therefore that δ 18 O values in plant cellulose can be used as a proxy for VPD in tropical ecosystems. As VPD is an essential variable that determines the biogeochemical dynamics of ecosystems, our study has applications in ecological-, climate-, or forensic-sciences.

Summary 1. Seasonal and inter-annual variability in ecosystem carbon dioxide (CO2) exchange is attributed to numerous climate drivers. However, climate effects on metabolism often override ecological functions. This study seeks insight into which biological and ecological processes influence temporal patterns of ecosystem productivity in natural ecosystems. 2. The specific objectives of this study are to (i) identify seasonal and inter-annual patterns of ecosystem-level photosynthesis in relation to climatic conditions, (ii) examine and compare seasonal and inter-annual variations in leaf traits for annual grasses and oak trees across multiple years, and (iii) explore interactions among leaf traits and ecosystem-level photosynthesis across multiple seasons and years. 3. We conducted this study in a woody savanna and open grassland in California, USA. Ecosystem-level photosynthetic rates of annual grasses (Agrass) and oak tree canopy (Acanopy) were deduced from eddy covariance measurements over a 7-year period (2001 and 2007). In conjunction, we sampled grass and oak leaves at weekly to monthly intervals and constructed a multiyear time series of leaf nitrogen concentration (N), leaf mass per unit area (LMA), leaf carbon concentration (C), and leaf carbon stable isotope discrimination (D). 4. Given the same grass community age or tree canopy age, inter-annual variations of the photosynthetic rates were up to 1–2 gC m )2 day )1 for annual grasses and oak trees while the two types of vegetation were exposed to different, wide ranges of inter-annual climate fluctuations: up to 5 � C in daily mean soil temperature, 15% in soil moisture, and 10 mol m )2 day )1 in photosynthetically active radiation. 5. While both grass and oak leaf traits varied seasonally and inter-annually, they experienced temporal patterns and seasonal peaks that were distinct from one another. Multi-year means of grass leaf N, C, D, and LMA were 2AE3%, 40AE8%, 22AE6& and 71AE 3gm )2 , respectively; multi-year means of oak leaf N, C, D, and LMA were 1AE9%, 45AE1%, 20AE5& and 132 g m )2 , respectively. 6. Based on the analysis of variance, seasonal and inter-annual terms were associated with Agrass or Acanopy up to 90% or 81%. On the other hand, variations in leaf N, LMA, C, D, and their interactions could statistically explain about 53% and 26% of variations in Agrass and Acanopy, respectively. 7. We discussed possible biological and ecological processes involved in regulating seasonal and inter-annual variability in ecosystem-level photosynthesis. Clearly, seasonal and inter-annual variation in ecosystem photosynthesis was strongly associated with the dynamics of leaf traits.

Biogeographical, physiological, and paleoecological evidence suggests that the coast redwood [ Sequoia sempervirens (D. Don) Endl.] is closely associated with the presence of summer marine fog along the Pacific coast of California. Here we present a novel record of summer fog frequency in the coast redwood region upon the basis of direct hourly measurements of cloud ceiling heights from 1951 to 2008. Our analysis shows that coastal summer fog frequency is a remarkably integrative measure of United States Pacific coastal climate, with strong statistical connections to the wind-driven upwelling system of the California Current and the broad ocean temperature pattern known as the Pacific Decadal Oscillation. By using a long-term index of daily maximum land temperatures, we infer a 33% reduction in fog frequency since the early 20th century. We present tree physiological data suggesting that coast redwood and other ecosystems along the United States west coast may be increasingly drought stressed under a summer climate of reduced fog frequency and greater evaporative demand.

The stable oxygen isotope ratio (delta(18)O) of plant material has been shown to contain essential information on water and carbon fluxes at the plant and ecosystem scales. However, the effective path length (L(m)), a parameter introduced to leaf-water models still requires a comprehensive biological characterization to allow interpretation of delta(18)O values in plant material with confidence. Here, we tested the variability of L(m) across and within three species that developed leaves in environments with different relative humidity. We also tested whether the L(m) of fully developed leaves is affected by short-term fluctuations in relative humidity. We determined that significant differences in L(m) exist among Phaseolus vulgaris, Rizinus communis and Helianthus annuus. Within a given species, however, L(m) values did not differ significantly among individuals. These findings indicate that L(m) is species specific and a relatively constant parameter and that L(m) will not obscure the interpretation of delta(18)O values in plant material of a given species. We urge caution, however, because values for L(m) are derived from fitting leaf-water models to measured values of delta(18)O, so care must be taken in assigning a 'cause' to values of L(m) as they likely capture a combination of different biological leaf properties.

Characterizing the use of carbon (C) reserves in trees is important for understanding regional and global C cycles, stress responses, asynchrony between photosynthetic activity and growth demand, and isotopic exchanges in studies of tree physiology and ecosystem C cycling. Using an inadvertent, whole-ecosystem radiocarbon (14C) release in a temperate deciduous oak forest and numerical modeling, we estimated that the mean age of stored C used to grow both leaf buds and new roots is 0.7 years and about 55% of new-root growth annually comes from stored C. Therefore, the calculated mean age of C used to grow new-root tissue is ∼0.4 years. In short, new roots contain a lot of stored C but it is young in age. Additionally, the type of structure used to model stored C input is important. Model structures that did not include storage, or that assumed stored and new C mixed well (within root or shoot tissues) before being used for root growth, did not fit the data nearly as well as when a distinct storage pool was used. Consistent with these whole-ecosystem labeling results, the mean age of C in new-root tissues determined using ‘bomb-14C’ in three additional forest sites in North America and Europe (one deciduous, two coniferous) was less than 1–2 years. The effect of stored reserves on estimated ages of fine roots is unlikely to be large in most natural abundance isotope studies. However, models of root C dynamics should take stored reserves into account, particularly for pulse-labeling studies and fast-cycling roots (

Fog is a seasonally variable hydrologic input to coast redwood ( Sequoia sempervirens, D. Don) ecosystems with a stable isotopic composition (δ 18O, δ2H) that is distinct from rainfall. The intra-annual variation in tree ring cellulose δ18O of coast redwood was measured to determine what portion of the tree ring should be sampled to best target fog signals. Ten years of rings were subdivided into ten equal units for three trees at two sites in northern California. Intra-annual variation in cellulose δ18O was as high as 3‰ with a consistent pattern of enriched (in 18O) values from the annual ring boundary and depleted values from the central portion of the ring. These patterns show a strong coherence between replicate trees at the same site as well as significant correlations between individuals at sites over 250 km apart. Intra-annual variation in stable carbon isotope ratios (δ13C) was as high as 2‰ and produced a pattern with depleted (in 13C) values from the ring boundary and enriched values from the central portion of the ring. Although these δ13C patterns were not as strongly correlated between replicate trees as the patterns in δ18O variation, they did produce significant correlations with the variation in fog frequencies recorded at local airports. This study highlights the importance of quantifying intra-annual variation in tree ring stable isotope signals as a guide to further investigations on historic variability from long chronologies especially if the signal of interest is seasonally variable (eg, fog).

Stable oxygen isotope ratios (delta18O) have become a valuable tool in the plant and ecosystem sciences. The interpretation of delta18O values in plant material is, however, still complicated owing to the complex interactions among factors that influence leaf water enrichment. This study investigated the interplay among environmental parameters, leaf physiological properties and leaf water relations as drivers of the isotopic enrichment of leaf water across 17 Eucalyptus species growing in a common garden. We observed large differences in maximum daily leaf water delta18O across the 17 species. By fitting different leaf water models to these empirical data, we determined that differences in leaf water delta18O across species are largely explained by variation in the Péclet effect across species. Our analyses also revealed that species-specific differences in transpiration do not explain the observed differences in delta18O while the unconstrained fitting parameter 'effective path length' (L) was highly correlated with delta18O. None of the leaf morphological or leaf water related parameters we quantified in this study correlated with the L values we determined even though L was typically interpreted as a leaf morphological/anatomical property. A sensitivity analysis supported the importance of L for explaining the variability in leaf water delta18O across different species. Our investigation highlighted the importance of future studies to quantify the leaf properties that influence L. Obtaining such information will significantly improve our understanding of what ultimately determines the delta18O values of leaf water across different plant species.

About half of the Amazon rainforest is subject to seasonal droughts of 3 months or more. Despite this drought, several studies have shown that these forests, under a strongly seasonal climate, do not exhibit significant water stress during the dry season. In addition to deep soil water uptake, another contributing explanation for the absence of plant water stress during drought is the process of hydraulic redistribution; the nocturnal transfer of water by roots from moist to dry regions of the soil profile. Here, we present data on patterns of soil moisture and sap flow in roots of three dimorphic-rooted species in the Tapajós Forest, Amazônia, which demonstrate both upward (hydraulic lift) and downward hydraulic redistribution. We measured sap flow in lateral and tap roots of our three study species over a 2-year period using the heat ratio method, a sap-flow technique that allows bi-directional measurement of water flow. On certain nights during the dry season, reverse or acropetal flow (i.e.,in the direction of the soil) in the lateral roots and positive or basipetal sap flow (toward the plant) in the tap roots of Coussarea racemosa (caferana), Manilkara huberi (maçaranduba) and Protium robustum (breu) were observed, a pattern consistent with upward hydraulic redistribution (hydraulic lift). With the onset of heavy rains, this pattern reversed, with continuous night-time acropetal sap flow in the tap root and basipetal sap flow in lateral roots, indicating water movement from wet top soil to dry deeper soils (downward hydraulic redistribution). Both patterns were present in trees within a rainfall exclusion plot (Seca Floresta) and to a more limited extent in the control plot. Although hydraulic redistribution has traditionally been associated with arid or strongly seasonal environments, our findings now suggest that it is important in ameliorating water stress and improving rain infiltration in Amazonian rainforests. This has broad implications for understanding and modeling ecosystem process and forest function in this important biome.

This study examines the effect of four tree species on nitrogen (N) retention within forested catchments of the Catskill Mountains, New York (NY). We conducted a 300-day 15N field tracer experiment to determine how N moves through soil, microbial, and plant pools under different tree species and fertilization regimes. Samples were collected from single-species plots of American beech (Fagus grandifolia Ehrh.), eastern hemlock (Tsuga canadensis L.), red oak (Quercus rubra L.), and sugar maple (Acer saccharum Marsh). Using paired plots we compared the effects of ambient levels of N inputs (11 kg N/ha/y) to additions of 50 kg N/ha/y that began 1.5 years prior to and continued throughout this experiment. Total plot 15N recovery (litter layer, organic and mineral soil to 12 cm, fine roots, and aboveground biomass) did not vary significantly among tree species, but the distribution of sinks for 15N within the forest ecosystem did vary. Recovery in the forest floor was significantly lower in sugar maple stands compared to the other species. 15Nitrogen recovery was 22% lower in the fertilized plots compared to the ambient plots and red oak stands had the largest drop in 15N recovery as a result of N fertilization. Aboveground biomass became a significantly greater 15N sink with fertilization, although it retained less than 1% of the tracer addition. These results indicate that different forest types vary in the amount of N retention in the forest floor, and that forest N retention may change depending upon N inputs.

Watersheds of the Catskill Mountains, New York have marked differences in nitrogen (N) dynamics among dominant tree species stands. Our objectives were to study how tree species vary in N uptake to better understand the basis for the observed variation in these forested watersheds. We conducted a 15N tracer greenhouse study to determine NH4 + and NO3 − uptake of American beech (Fagus grandifolia Ehrh.), eastern hemlock (Tsuga Canadensis L.), red oak (Quercus rubra L.) and sugar maple (Acer saccharum Marsh.) seedlings. Seedlings and their native soil were collected in November 1997, over-wintered and allowed to break dormancy in spring 1998. Half of the seedlings of each tree species received 15NH4-NO3 to examine NH4 + uptake and the other half received NH4-15NO3 to examine NO3 − uptake. Plants were harvested 4 days following 15N addition. Tree species varied in their preference for NH4 + and NO3 −. Sugar maple and eastern hemlock seedlings took up more NH4 + than NO3 − per unit plant biomass, while beech was the only species to take up more NO3 − than NH4 +. Red oak took up more NH4 + than NO3 − into roots, stems and leaves, but the difference between the two forms of N was not statistically significant. These results demonstrate that tree species of the Catskill Mountains vary in their capacity to take up NH4 + and NO3 −. Coupled with stand-level studies of N dynamics, this variation can help explain some of the patterns of forested watershed N retention and loss in the Catskill Mountains shown in our field investigations (Templer, 2001; Templer et al., in press Ecosystems).

Recent studies suggest that savanna trees in semi-arid areas can increase understorey plant production. We hypothesized that one of the mechanisms that explains the facilitation between trees and grasses in East African savannas is hydraulic lift (HL). HL in large Acacia tortilis trees was studied during the first 3 months of the dry season during a relatively wet year (1998) and a very dry year (2000). In 1998, we found distinct diel fluctuation in soil water potential (psi(s)), with increasing values during the night and decreasing again the following day. These fluctuations in psi(s )are consistent with other observations of HL and in A. tortilis were found up to 10 m from the tree. In 2000, during a severe drought, psi(s) measurements indicated that HL was largely absent. The finding that HL occurred in wetter years and not in drier years was supported by data obtained on the delta(18)O values in soil, rain and groundwater. The delta(18)O of water extracted from the xylem water of grasses indicated that when they grew near trees they had values similar to those of groundwater. This could be because they either (1) use water from deeper soil layers or (2) use hydraulically lifted water provided by the tree; this was not seen in the same grass species growing outside tree canopies. While our data indicate that HL indeed occurs under Acacia trees, it is also true that psi(s) was consistently lower under trees when compared to outside tree canopies. We believe that this is because tree-grass mixtures take up more water from the upper soil layers than is exuded by the tree each night. This limits the beneficial effect of HL for understorey grasses and suggests that in savannas both facilitation via HL and competition are active processes. The importance of each process may depend upon how wet or dry that particular site or year is.

▪ Abstract The use of stable isotope techniques in plant ecological research has grown steadily during the past two decades. This trend will continue as investigators realize that stable isotopes can serve as valuable nonradioactive tracers and nondestructive integrators of how plants today and in the past have interacted with and responded to their abiotic and biotic environments. At the center of nearly all plant ecological research which has made use of stable isotope methods are the notions of interactions and the resources that mediate or influence them. Our review, therefore, highlights recent advances in plant ecology that have embraced these notions, particularly at different spatial and temporal scales. Specifically, we review how isotope measurements associated with the critical plant resources carbon, water, and nitrogen have helped deepen our understanding of plant-resource acquisition, plant interactions with other organisms, and the role of plants in ecosystem studies. Where possible we also introduce how stable isotope information has provided insights into plant ecological research being done in a paleontological context. Progress in our understanding of plants in natural environments has shown that the future of plant ecological research will continue to see some of its greatest advances when stable isotope methods are applied.

Many species have the ability to resprout vegetatively after a substantial loss of biomass induced by environmental stress, including drought. Many of the regions characterised by ecosystems where resprouting is common are projected to experience more frequent and intense drought during the 21st Century. However, in assessments of ecosystem response to drought disturbance there has been scant consideration of the resilience and post-drought recovery of resprouting species. Systematic differences in hydraulic and allocation traits suggest that resprouting species are more resilient to drought-stress than nonresprouting species. Evidence suggests that ecosystems dominated by resprouters recover from disturbance more quickly than ecosystems dominated by nonresprouters. The ability of resprouters to avoid mortality and withstand drought, coupled with their ability to recover rapidly, suggests that the impact of increased drought stress in ecosystems dominated by these species may be small. The strategy of resprouting needs to be modelled explicitly to improve estimates of future climate-change impacts on the carbon cycle, but this will require several important knowledge gaps to be filled before resprouting can be properly implemented.

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

Although clouds are the most recognisable and defining feature of tropical montane cloud forests, little research has focussed on how clouds affect plant functioning. We used satellite and ground-based observations to study cloud and leaf wetting patterns in contrasting tropical montane and pre-montane cloud forests. We then studied the consequences of leaf wetting for the direct uptake of water accumulated on leaf surfaces into the leaves themselves. During the dry season, the montane forest experienced higher precipitation, cloud cover and leaf wetting events of longer duration than the pre-montane forest. Leaf wetting events resulted in foliar water uptake in all species studied. The capacity for foliar water uptake differed significantly between the montane and pre-montane forest plant communities, as well as among species within a forest. Our results indicate that foliar water uptake is common in these forest plants and improves plant water status during the dry season.

Assessing the ecological importance of clouds has substantial implications for our basic understanding of ecosystems and for predicting how they will respond to a changing climate. This study was conducted in a coastal Bishop pine forest ecosystem that experiences regular cycles of stratus cloud cover and inundation in summer. Our objective was to understand how these clouds impact ecosystem metabolism by contrasting two sites along a gradient of summer stratus cover. The site that was under cloud cover ~15% more of the summer daytime hours had lower air temperatures and evaporation rates, higher soil moisture content, and received more frequent fog drip inputs than the site with less cloud cover. These cloud-driven differences in environmental conditions translated into large differences in plant and microbial activity. Pine trees at the site with greater cloud cover exhibited less water stress in summer, larger basal area growth, and greater rates of sap velocity. The difference in basal area growth between the two sites was largely due to summer growth. Microbial metabolism was highly responsive to fog drip, illustrated by an observed ~3-fold increase in microbial biomass C with increasing summer fog drip. In addition, the site with more cloud cover had greater total soil respiration and a larger fractional contribution from heterotrophic sources. We conclude that clouds are important to the ecological functioning of these coastal forests, providing summer shading and cooling that relieve pine and microbial drought stress as well as regular moisture inputs that elevate plant and microbial metabolism. These findings are important for understanding how these and other seasonally dry coastal ecosystems will respond to predicted changes in stratus cover, rainfall, and temperature.

Summary Variation in the stable carbon and oxygen isotope composition (δ13C, Δ18O) of co-occurring plant species may reflect the functional diversity of water use strategies present in natural plant communities. We investigated the patterns of water use among 10 coexisting plant species representing diverse taxonomic groups and life forms in semiarid southeast Spain by measuring their leaf δ13C and Δ18O, the oxygen isotope ratio of stem water and leaf gas exchange rates. Across species, Δ18O was tightly negatively correlated with stomatal conductance (gs), whereas δ13C was positively correlated with intrinsic water use efficiency (WUEi). Broad interspecific variation in Δ18O, δ13C and WUEi was largely determined by differences in gs, as indicated by a strong positive correlation between leaf δ13C and Δ18O across species The 10 co-occurring species segregated along a continuous ecophysiological gradient defined by their leaf δ13C and Δ18O, thus revealing a wide spectrum of stomatal regulation intensity and contrasting water use strategies ranging from ‘profligate/opportunistic’ (high gs, low WUEi) to ‘conservative’ (low gs, high WUEi). Coexisting species maintained their relative isotopic rankings in 2 yr with contrasting rainfall, suggesting the existence of species-specific ‘isotopic niches’ that reflect ecophysiological niche segregation in dryland plant communities.

Moisture inputs drive soil respiration (SR) dynamics in semi-arid and arid ecosystems. However, determining the contributions of root and microbial respiration to SR, and their separate temporal responses to periodic drought and water pulses, remains poorly understood. This study was conducted in a pine forest ecosystem with a Mediterranean-type climate that receives seasonally varying precipitation inputs from both rainfall (in the winter) and fog-drip (primarily in the summer). We used automated SR measurements, radiocarbon SR source partitioning, and a water addition experiment to understand how SR, and its separate root and microbial sources, respond to seasonal and episodic changes in moisture. Seasonal changes in SR were driven by surface soil water content and large changes in root respiration contributions. Superimposed on these seasonal patterns were episodic pulses of precipitation that determined the short-term SR patterns. Warm season precipitation pulses derived from fog-drip, and rainfall following extended dry periods, stimulated the largest SR responses. Microbial respiration dominated these SR responses, increasing within hours, whereas root respiration responded more slowly over days. We conclude that root and microbial respiration sources respond differently in timing and magnitude to both seasonal and episodic moisture inputs. These findings have important implications for the mechanistic representation of SR in models and the response of dry ecosystems to changes in precipitation patterns.

 Premise of the study : Fog provides a critical water resource to plants around the world. In the redwood forest ecosystem of northern California, plants depend on fog absorbed through foliar uptake to stay hydrated during the rainless summer. In this study, we identifi ed regions within the redwood ecosystem where the fern Polystichum munitum canopy most effectively absorbs fog drip that reaches the forest fl oor.  Methods : We measured the foliar uptake capacity of P. munitum fronds at seven sites along 700 km of the redwood forest ecosystem. We quantifi ed the canopy cover of P. munitum at each site and estimated how much water the fern canopy can acquire aboveground through fog interception and absorption.  Key results : Throughout the ecosystem, nocturnal foliar uptake increased the leaf water content of P. munitum by 7.2%, and we estimated that the P. munitum canopy can absorb 5 ± 3% (mean ± SE) of intercepted fog precipitation. Strikingly, P. munitum had the highest foliar uptake capacity in the center of the ecosystem and may absorb 10% more of the fog its canopy intercepts in this region relative to other regions studied. Conversely, P. munitum had no foliar uptake capacity in the southern end of the ecosystem.  Conclusions : This study shows the fi rst evidence that foliar uptake varies within species at the landscape scale. Our fi ndings suggest that the P. munitum at the southern tip of the redwood ecosystem may suffer most from low summertime water availability because it had no potential to acquire fog as an aboveground water subsidy.

List of contributors Preface SECTION I: THE WHOLE-LIFESPAN PERSPECTIVE ON INTEGRATION OF STRUCTURE AND FUNCTION 1. A Lifespan Perspective on Integrating Structure and Function in Trees Thomas M. Hinckley, Barbara Lachenbruch, Frederick C. Meinzer, and Todd E. Dawson SECTION II: REPRODUCTION AND REGENERATION 2. Age-related Changes in Tree Growth and Functional Biology: the Role of Reproduction Sean C. Thomas 3. The Earliest Stages of Tree Growth: Development, Physiology and Impacts of Climate Change Daniel Johnson, Katherine A. McCulloh, and Keith Reinhardt SECTION III: INTEGRATION OF STRUCTURE AND FUNCTION ACROSS ONTOGENY 4, Regulation of Ontogeny in Temperate Conifers Michael E. Day and Michael S. Greenwood 5. Radial Variation in Wood Structure and Function in Woody Plants, and Hypotheses for its Occurrence Barbara Lachenbruch, John R. Moore, and Robert Evans 6. Size-related Changes in Tree Proportions and Their Potential Influence on the Course of Height Growth David A. King 7. Influence of Tree Ontogeny on Plant-Herbivore Interactions Karina Boege, Kasey E. Barton, and Rodolfo Dirzo 8. How do Changes in Leaf/shoot Morphology and Crown Architecture Affect Growth and Physiological Function of Large, old Trees? Hiroaki Ishii SECTION IV: PHOTOSYNTHETIC GAS EXCHANGE AND CARBON BALANCE 9. Tree size- and Age-Related Changes in Leaf Physiology and Their Influence on Carbon Gain Kathy Steppe, Ulo Niinemets, and Robert O. Teskey 10. Relationships Between Tree Height and Carbon Isotope Discrimination Nate G. McDowell, Barbara J. Bond, Lee T. Hill, Michael G. Ryan, and David Whitehead 11. Carbon Storage in Trees: Does Relative Carbon Supply Decrease With Tree Size? Anna Sala,Willa Fouts, and Gunter Hoch SECTION V: WATER RELATIONS AND TRANSPORT PROCESSES 12. Comparative Criteria for Models of the Fluid Transport System of Tall Trees Maurizio Mencuccini, Teemu Holtta, and Jordi Martinez-Vilalta 13. Hydraulic Capacitance: Biophysics and Functional Significance of Internal Water Sources in Relation to Tree Size Fabian G. Scholz, Nathan G. Phillips, Sandra J. Bucci, Frederick C. Meinzer, and Guillermo Goldstein 14. Size-Dependent Changes in Biophysical Control of Tree Growth: the Role of Turgor David R. Woodruff and Frederick C. Meinzer 15. Instrumental Approaches to Studying Tree Water Relations Along Gradients of Tree Size and Forest Age Jan Cermak and Nadezhda Nadezhdina SECTION VI: SCALING FROM INDIVIDUAL TRAITS TO STANDS 16. Ecophysiology of Secondary Succession in Tropical Moist Forest: Scaling From Individual Traits to Whole-Plant Performance Niels Anten and N. Galia Selaya 17. Age-Dependent Changes in Environmental Influences on Tree Growth and Their Implications for Forest Responses to Climate Change Steven L. Voelker 18. Consequences of Stand age and Species' Functional Trait Changes on Ecosystem Water use of Forests Brent Ewers, D. Scott Mackay, and Ben Bond-Lamberty

Trees are spectacular organisms that can accumulate a large amount of biomass, live for millennia, grow in stressful environments, and have global importance to organisms and human society as a result of their roles in contributing to diverse ecosystem services. They survive in an enormous range of environments with disturbances of various frequencies and severities, and by their very nature of being long-lived and sessile, they must possess the ability to change. This chapter provides a context for the chapters that follow, by describing a history of the investigations and of several important technological innovations that have enabled research to progress, and some of the foundations of water relations which underpin many of the later chapters. The chapter then discusses seven recurrent themes brought out by the diverse chapter authors: microclimate and other abiotic forces that change with tree and stand age; the changing structure and function employed as trees grow; what we can learn from scaling —which scaling laws pertain, when, and what is the biology or physics embedded in these laws; the tradeoffs that occur as structures and functions change through growth and development; the causes and correlates of the decline in productivity of trees and stands with age; the extent to which changes are plastic; and the meaning of whether changes occur relative to age versus size. The chapter concludes with a discussion on the emerging issues and questions raised in the chapters of this book. The overall aim is to give insights into the whole tree, whole lifespan perspective of the structural and functional strategies trees employ to persist in the face of internal and external factors that change as trees grow and age. This knowledge can be used to develop management approaches to manipulate trees and stands to provide a wide range of ecosystem goods and services.

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Evaluations of plant water use in ecosystems around the world reveal a shared capacity by many dif- ferent species to absorb rain, dew, or fog water directly into their leaves or plant crowns. This mode of water uptake provides an important water subsidy that relieves foliar water stress. Our study provides the first comparative evaluation of foliar uptake capacity among the dominant plant taxa from the coast redwood ecosystem of California where crown-wetting events by summertime fog frequently occur during an otherwise drought-prone season. Previous research demonstrated that the dominant overstory tree species, Sequoia sempervirens, takes up fog water by both its roots (via drip from the crown to the soil) and directly through its leaf surfaces. The present study adds to these early findings and shows that 80% of the dominant species from the redwood forest exhibit this foliar uptake water acquisition strategy. The plants studied include canopy trees, understory ferns, and shrubs. Our results also show that foliar uptake provides direct hydration to leaves, increasing leaf water content by 2-11%. In addition, 60% of redwood forest species investigated demonstrate noc- turnal stomatal conductance to water vapor. Such findings indicate that even species unable to absorb water directly into their foliage may still receive indirect benefits from nocturnal leaf wetting through suppressed transpiration. For these species, leaf-wetting events enhance the efficacy of nighttime re-equilibration with available soil water and therefore also increase pre-dawn leaf water potentials.

13 páginas, 3 tablas, 7 figuras., Ratios of nitrogen (N) isotopes in leaves could elucidate underlying patterns of N cycling across ecological gradients. To better understand global-scale patterns of N cycling, we compiled data on foliar N isotope ratios (δ15N), foliar N concentrations, mycorrhizal type and climate for over 11 000 plants worldwide. Arbuscular mycorrhizal, ectomycorrhizal, and ericoid mycorrhizal plants were depleted in foliar δ15N by 2‰, 3.2‰, 5.9‰, respectively, relative to nonmycorrhizal plants. Foliar δ15N increased with decreasing mean annual precipitation and with increasing mean annual temperature (MAT) across sites with MAT ≥ −0.5°C, but was invariant with MAT across sites with MAT < −0.5°C. In independent landscape-level to regional-level studies, foliar δ15N increased with increasing N availability; at the global scale, foliar δ15N increased with increasing foliar N concentrations and decreasing foliar phosphorus (P) concentrations. Together, these results suggest that warm, dry ecosystems have the highest N availability, while plants with high N concentrations, on average, occupy sites with higher N availability than plants with low N concentrations. Global-scale comparisons of other components of the N cycle are still required for better mechanistic understanding of the determinants of variation in foliar δ15N and ultimately global patterns in N cycling., J.M.C. was supported by the Andrew W. Mellon Foundation and A.E. was supported by the Henry Luce Foundation.

Publisher Summary The stable isotope investigations have grown steadily in the past 50 years or so and have become a cornerstone for many ecological investigations since the 1980s. The understanding of the many processes that lead to isotopic variation in natural environments is now often well understood and is improving each year. The challenge that is now being confronted with, is to extend what is known about these processes and the products that result from them, to see how changes have been ‘‘recorded’’ and how these changes can be interpreted. This requires a different approach that many past ecological investigations have not taken and one that forces ecologists to find materials such as carbonates, feathers, hard tissues, hair, tree-ring cellulose, and the many chemical compounds that compose them, so that they can be used to trace and therefore record change. By linking known processes to these changes, the growing field of ‘‘ecological forensics'’ that has emerged, is now documenting changes in many novel and powerful ways.Through innovative stable isotope methods and their novel application, the manner and magnitude of ecological transformations are now being documented across a wide range of natural systems. The environmental and ecological changes will continue and in many instances, will accelerate. Applying stable isotope analyses in these ways can lead to solutions that will help mitigate future ecosystem damage and environmental degradation.

Variation in flower size is an important aspect of a plant's life history, yet few studies have shown how flower size varies with environmental conditions and to what extent foliar responses to the environment are correlated with flower size. The objectives of this study were to (1) develop a theoretical framework for linking flower size and leaf size to their costs and benefits, as assessed using foliar stable carbon isotope ratio (delta(13)C) under varying degrees of water limitation, and then (2) examine how variation in flower size within and among species growing along a naturally occurring moisture availability gradient correlates with variation in delta(13)C and leaf size. Five plant species were examined at three sites in Oregon. Intra- and inter-specific patterns of flower size in relation to moisture availability were the same: the ratios of the area of flower display to total leaf area and of individual flower area to leaf area were greater at sites with more soil moisture compared to those sites with less soil moisture. The increase in flower area per unit increase in leaf area was greater at sites with more soil moisture than at sites where water deficit can occur. Values of delta(13)C, an index of water-use efficiency, were greater for plants with larger floral size. The patterns we observed generalize across species, irrespective of overall plant morphology or pollination system. These correlations between flower size, moisture availability, and delta(13)C suggest that water loss from flowers can influence leaf responses to the environment, which in turn may indirectly mediate an effect on flower size.

Recent studies have shown that natural abundance stable carbon isotopes can be used to partition measurements of net ecosystem CO 2 exchange into gross fluxes of photosynthesis and respiration. This chapter explores the related application of the natural abundance stable carbon isotope ratio of respired CO 2 to partitioning components of ecosystem respiration. Although, the exploitation of the natural variation in stable isotope abundance in respired CO 2 holds the most promise for widespread application among existing measurement networks (e.g., BASIN, FLUXNET, SIBAE) and for comparative studies focused on ecosystem C balance, the reliance on natural variation may also have limitations from the standpoint that such variation is typically small. Recent evidence suggests, however, that variation can be large and significant such that measurable differences among respiration signatures can allow partitioning of ecosystem respiration and therefore inform ecosystem C-balance studies.

Understanding the responses of riparian trees to water availability is critical for predicting the effects of changes in precipitation on riparian ecosystems. Dioecious Acer negundo L. (box elder) is a common riparian tree that is highly sensitive to water stress. Earlier studies indicated that the genders of A. negundo respond differently to water availability, with males being more conservative in their water use than females. To assess the potential effects of changes in precipitation on the sex ratio of riparian trees, we extended earlier studies of A. negundo by analyzing responses of male and female genotypes to interannual differences in water availability in a common garden. We measured growth of tree rings and used stable carbon isotope analysis of tree ring alpha-cellulose to integrate physiological responses to annual water treatments. During dry years, male and female trees exhibited similar growth and physiological responses. However, during wet years, females exhibited higher growth rates and lower carbon isotope ratios (indicating less conservative water use) than did males. Furthermore, we found that male trees exhibited similar stomatal behavior (inferred from integrated carbon isotope ratios) whether years were wet or dry, whereas females did not exhibit a consistent response to changes in water availability. We predict that with increasing precipitation and soil water availability, the representation of females will be favored because of shifts in the competitive interactions of the genders. Such changes may affect the reproductive output of these riparian trees and may influence overall water flux from riparian ecosystems. In addition, this study demonstrates the utility of carbon isotope analysis for assessing long-term responses of tree populations to shifts in water availability.