Identifying groundwater‐fed climate refugia in remote arid regions with citizen science and isotope hydrology

1. Groundwater and surface water‐fed systems act as biodiversity hotspots and ecological refuges and evolutionary refugia in arid regions. Groundwater‐dominated systems are sustained by underground aquifers that are recharged by rain that has fallen in the distant past, while surface water‐dominated systems are fed by recent local rain or floods. Some waterbodies are fed by a mixture of these sources. Perennial, groundwater‐dominated systems will act as refuges and refugia under future rainfall declines associated with global warming. We sought to identify climate refugia, based on groundwater dominance, by using isotope hydrology to characterise water samples collected by citizen scientists across arid central Australia.
2. There is a linear relationship between hydrogen isotopes (²H/¹H, δ²H) and oxygen isotopes (¹⁸O/¹⁶O, δ¹⁸O) in rainfall. This relationship is known as the meteoric water line (MWL). By comparing our samples with the Australian MWL, and developing a local evaporation line, we were able to test the hypotheses that groundwater‐dominated systems will follow the Australian MWL while temporary systems follow the local evaporation line, and, accordingly, distinguish between groundwater and surface water‐dominated systems. The isotopic composition of samples collected over a 36‐month period was determined using isotope ratio infrared spectrometry. The electrical conductivity of each sample was recorded to determine where freshwater is available for biota within this arid region.
3. Over 240 water samples were collected from 62 waterbodies and seven bores (groundwater wells) spanning an area of more than 250,000 km². Approximately 75% of the samples were collected by citizen scientists and 25% by research scientists.
4. Twenty groundwater‐dominated waterbodies, characterised by a small range of δ²H and δ¹⁸O values (c. ?55 to ?20‰ and c. ?9 to ?3‰, respectively) clustered around the long‐term mean composition of rainfall (δ²H = ?37.5‰, δ¹⁸O = ?6.4‰), were identified as future evolutionary refugia. These sites are likely to contain water through the most severe of droughts and will be critically important for the persistence of water‐dependent species.
5. Based on their isotopic composition, we identified 45 waterbodies (rockholes/waterholes) as temporary or ephemeral (δ²H c. ?40 to ?100‰ and δ¹⁸O c. ?4 to +25‰), that is, with no evidence of groundwater inflow. These, together with waterbodies supported by a mix of groundwater and surface water, can act as stepping stones and form part of the aquatic mosaic that is critical to supporting species in arid regions. Over two‐thirds of the waterholes sampled were very fresh (electrical conductivity <0.8 mS/cm), indicating that they provide the freshwater needed to support much of the regional aquatic and terrestrial fauna.
6. All evolutionary refugia are located within protected areas (i.e. national parks or Indigenous Protected Areas), but some are subject to the impacts of feral animal species and invasive plants. Our findings indicate where control programmes and restoration actions can be prioritised to support biodiversity conservation and climate change adaptation. Our approach, combining citizen science and isotope hydrology, can be used to identify future refugia in other remote and arid regions where water scarcity is likely to increase under global climate change.

     

Tree age,site and climate controls on tree ring cellulose δ18O: A case study on oak trees from south-western France

A main concern of dendroclimatic reconstruction is to distinguish in the tree ring proxy the influence of the climate variables of interest from other controlling factors. In order to investigate age, site and climate controls on tree ring width and cellulose δ¹⁸O, measurements have been performed in nearby groups of young (145 years old) and older (310–405 years old) oak trees in south-western France, covering the period 1860–2010.Within a given site, inter-tree deviations are small, pointing to a common climatic signal. Despite a similar inter-annual variability, the average level of cellulose δ¹⁸O in the young tree group is ∼0.8‰ higher than in the old trees. Such offsets might be caused by different soil properties and differences in the fraction of the source water used by trees from different depths. The δ¹⁸O of water in the top soil layer is directly related to the current growing season precipitation, while deeper water can have a lower and more constant δ¹⁸O. Local cave drip waters at 10 m depth indeed show a constant isotopic composition, which corresponds to pluri-annual mean precipitation.A 2‰ increasing trend is observed in cellulose δ¹⁸O of young trees in the first 30 years of growth, during a period when no trend is visible in older trees. This increase can be quantitatively explained by humidity gradients under the forest canopy, and a changing microclimate around the crown as trees grow higher.While relationships between tree ring width and climate appear complex, the isotopic composition of cellulose is strongly correlated with summer maximum temperature, relative humidity and evapotranspiration (r ≈ 0.70). Weaker correlations (r ≈ 0.40) are identified with precipitation δ¹⁸O from a 15-year long local record and from the REMOiso model output. These results imply that leaf water enrichment has a stronger control on the inter-annual variability of cellulose δ¹⁸O than the δ¹⁸O of precipitation.This study demonstrates the suitability of oak tree ring cellulose δ¹⁸O for reconstructing past summer climate variability in south-western France, provided that the sampling and pooling strategy accounts for the fact that trees from different sites and of different age can introduce non-climatic signals.     

Exploring climate forcing of growth depression in subfossil South Swedish bog pines using stable isotopes

Comparison between growth variability, based on ring-width (RW) analysis, and moisture-sensitive signals in tree-ring carbon and oxygen stable-isotope composition provides increased understanding of how climate and hydrology influenced bog pines (Pinus sylvestris L.) at two sites in southern Sweden during the mid- and late Holocene. Tree-ring sequences from two subfossil trees collected at raised bogs having different hydrology and catchment size were analyzed to probe the stable-isotope signals associated with two bog-wide episodes of growth depression, one during the Holocene Thermal Maximum and the other during the Neoglacial Transition. The occurrence of lower whole-wood δ¹³C and cellulose δ¹³C and δ¹⁸O values immediately prior to the onset of growth depression in both trees, suggesting increased atmospheric relative humidity, is consistent with the notion that excessive effective moisture impeded tree growth. Correlation analysis indicates that the growth response lagged about three years behind the decline in δ¹³C and δ¹⁸O values in each tree, possibly reflecting relatively slow rise in the local water table in response to wetter climate.     

Increasing carbon discrimination rates and depth of water uptake favor the growth of Mediterranean evergreen trees in the ecotone with temperate deciduous forests

Tree populations at the low‐altitudinal or ‐latitudinal limits of species' distributional ranges are predicted to retreat toward higher altitudes and latitudes to track the ongoing changes in climate. Studies have focused on the climatic sensitivity of the retreating species, whereas little is known about the potential replacements. Competition between tree species in forest ecotones will likely be strongly influenced by the ecophysiological responses to heat and drought. We used tree‐ring widths and δ¹³C and δ¹⁸O chronologies to compare the growth rates and long‐term ecophysiological responses to climate in the temperate‐Mediterranean ecotone formed by the deciduous Fagus sylvatica and the evergreen Quercus ilex at the low altitudinal and southern latitudinal limit of F. sylvatica (NE Iberian Peninsula). F. sylvatica growth rates were similar to those of other southern populations and were surprisingly not higher than those of Q. ilex, which were an order of magnitude higher than those in nearby drier sites. Higher Q. ilex growth rates were associated with high temperatures, which have increased carbon discrimination rates in the last 25 years. In contrast, stomatal regulation in F. sylvatica was proportional to the increase in atmospheric CO₂. Tree‐ring δ¹⁸O for both species were mostly correlated with δ¹⁸O in the source water. In contrast to many previous studies, relative humidity was not negatively correlated with tree‐ring δ¹⁸O but had a positive effect on Q. ilex tree‐ring δ¹⁸O. Furthermore, tree‐ring δ¹⁸O decreased in Q. ilex over time. The sensitivity of Q. ilex to climate likely reflects the uptake of deep water that allowed it to benefit from the effect of CO₂ fertilization, in contrast to the water‐limited F. sylvatica. Consequently, Q. ilex is a strong competitor at sites currently dominated by F. sylvatica and could be favored by increasingly warmer conditions.     

Tree distribution on a steep environmental gradient in an arid savanna

Aim The structure of savannas ranges from scrub to woodland over broad geographical gradients. Here we examine the hypothesis that water availability is a major determinant of these structural differences by ascertaining the relationship between water availability and tree growth across a steep moisture gradient. Location The study site is a sub‐tropical savanna, with a mean annual precipitation of 400 mm, located in the Phugwane river basin, Kruger National Park, Limpopo Province, South Africa. Methods We determined plant moisture stress using xylem pressure potentials, stem growth using dendrometer bands, and estimated the water sources available to plants using stable hydrogen and oxygen isotope ratios. The primary objective was to understand tree growth relative to available water along an environmental gradient. Results We found that Philenoptera violacea trees growing close to the stream have lower water stress (least negative xylem pressure potentials) and higher cumulative growth than those growing away from the stream. The stem growth of P. violacea was characterized by steady incremental growth and could not be related to antecedent rainfall. Colophospermum mopane trees experienced higher water stress, yet trees growing adjacent to the stream achieved the highest cumulative growth rates over the study period. The growth of C. mopane could be clearly linked to antecedent rainfall, and most growth was achieved during short growth pulses that followed rainfall events. δ¹⁸O values become progressively more enriched in the heavier isotope with distance from the stream, suggesting that access to groundwater decreases with distance from the stream side. The isotopic data suggest that P. violacea has access to groundwater, but that C. mopane does not. Main conclusions Our results show that water stress increases with distance from the stream side as a result of reduced access to groundwater. Trees without access to deep water adopt an opportunistic growth strategy.     

Does resin tapping affect the tree-ring growth and climate sensitivity of the Chinese pine (Pinus tabuliformis) in the Loess Plateau,China?

Resin tapping could affect water and nutrient transport processes in Chinese pine trees, rendering them more vulnerable to extreme climatic events, such as drought, and affecting the ecological function of forests in semi-arid regions. This study evaluated how resin tapping affects the tree-ring growth and climate sensitivity of Chinese pine in the Loess Plateau. We compared tree-ring growth patterns between the tapped and untapped faces of tapped trees, and investigated tree-ring growth and its response to climate between tapped and untapped trees in a forest stand during the 1967–2017 period. Tapped trees showed asymmetrical growth patterns after resin tapping, with narrower rings near the tapped face and wider ones near the untapped face. Furthermore, tapped trees had inter-annual variations consistent with those of untapped trees except for the years 2000 and 2001, with significantly lower values following resin tapping, and tree-ring growth then returning to normal. The climate response analysis indicated that the tree-ring growth of both tapped and untapped trees was negatively affected by monthly mean temperatures during the early growing season (May to July) in the post-resin-tapping period. Furthermore, tree-ring growth in tapped trees also revealed significant correlation with water vapour deficit and the Palmer drought index, which indicates that tapped trees are more vulnerable to drought. Further studies based on stable isotopes (i.e. δ¹³C, δ¹⁸O, and δ¹⁵N) could improve our understanding of the physiological mechanisms that regulate the effects of resin tapping on tree-ring growth.     

Hominid environments at Hadar from paleosol studies in a framework of Ethiopian climate change

The amount and seasonal distribution of paleo-rainfall is a major concern of paleoanthropology because they determine the nature of the vegetation and the structure of the ecosystem, particularly in eastern Africa. The δ¹⁸O and δ¹³C of paleosol carbonates are quantitative proxies of these critical features of the paleoenvironment. The Afar region of Ethiopia lies between the African and Indian summer monsoons, and is prone to profound climate change. In the western Afar, the dominant paleoenvironment of the Hadar Formation during the late Pliocene was a major meandering river's distal low, flat floodplain, on which muds accreted that were continuously transformed into vegetated soils with Bk horizons rich in CaCO₃. The mean δ¹³C of paleosols throughout the Hadar Formation translates to an average vegetative cover across the extensive floodplain of about 30% of the C₄ grasses and 70% of unspecified C₃ plants. The character of the paleosols, such as the one at Locality 333, and their δ¹⁸O_Carbonate argue for a highly seasonal rainfall of about twice today's amount, implying that the C₃ plants were mostly sizeable trees and that the biome for Australopithecus afarensis was a grassy woodland. The amount of grasses abruptly increased in the lower Busidima Formation with its early Homo and artifacts to a more open grassy woodland of ca. 50% grasses. However, this transition in δ¹³C is not mirrored in the δ¹⁸O, which persists at a quite negative average value of −6.4‰ over the entire >2-Myr duration of both formations. This value for the carbonate means that the paleosoil water was a quite negative −4.1‰, a significant 5‰ more negative than our estimate of modern rain at Hadar. We put the negative δ¹⁸O of paleo-Hadar's rainfall into an isotopic framework of the dynamic history of climate change in sub-Saharan northern Africa. There have been two end-member climate regimes: (1) an earlier persistently pluvial Pliocene regime, with its strong summer monsoon, as registered in the Hadar Formation; and (2) the modern cyclical, mostly arid regime that began ca. 1 Myr ago, which has been punctuated by about ten cyclically predictable brief millennia-long pluvial episodes. The best known pluvial of the latter regime is the latest one, the African Humid Period (AHP), just 9.0-6.5 kyr ago, whose δ¹⁸O_Rainfall matches that for paleo-Hadar. The known climatological factors that brought on the AHP are probably the same ones that were persistently present for the Afar of the Pliocene. This dynamic rainfall history undoubtedly has influenced hominid occupation of the keystone Afar area at the gateway out of, and into, Africa.     

Plasticity of functional traits varies clinally along a rainfall gradient in Eucalyptus tricarpa

Widespread species often occur across a range of climatic conditions, through a combination of local genetic adaptations and phenotypic plasticity. Species with greater phenotypic plasticity are likely to be better positioned to cope with rapid anthropogenic climate changes, while those displaying strong local adaptations might benefit from translocations to assist the movement of adaptive genes as the climate changes. Eucalyptus tricarpa occurs across a climatic gradient in south‐eastern Australia, a region of increasing aridity, and we hypothesized that this species would display local adaptation to climate. We measured morphological and physiological traits reflecting climate responses in nine provenances from sites of 460 to 1040 mm annual rainfall, in their natural habitat and in common gardens near each end of the gradient. Local adaptation was evident in functional traits and differential growth rates in the common gardens. Some traits displayed complex combinations of plasticity and genetic divergence among provenances, including clinal variation in plasticity itself. Provenances from drier locations were more plastic in leaf thickness, whereas leaf size was more plastic in provenances from higher rainfall locations. Leaf density and stomatal physiology (as indicated by δ¹³C and δ¹⁸O) were highly and uniformly plastic. In addition to variation in mean trait values, genetic variation in trait plasticity may play a role in climate adaptation.     

The enigma of effective path length for 18O enrichment in leaf water of conifers

The Péclet correction is often used to predict leaf evaporative enrichment and requires an estimate of effective path length (L). Studies to estimate L in conifer needles have produced unexpected patterns based on Péclet theory and leaf anatomy. We exposed seedlings of six conifer species to different vapour pressure deficits (VPD) in controlled climate chambers to produce steady‐state leaf water enrichment (in ¹⁸O). We measured leaf gas exchange, stable oxygen isotopic composition (δ¹⁸O) of input and plant waters as well as leaf anatomical characteristics. Variation in bulk needle water δ¹⁸O was strongly related to VPD. Conifer needles had large amounts of water within the vascular strand that was potentially unenriched (up to 40%). Both standard Craig–Gordon and Péclet models failed to accurately predict conifer leaf water δ¹⁸O without taking into consideration the unenriched water in the vascular strand and variable L. Although L was linearly related to mesophyll thickness, large within‐species variation prevented the development of generalizations that could allow a broader use of the Péclet effect in predictive models. Our results point to the importance of within needle water pools and isolating mechanisms that need further investigation in order to integrate Péclet corrections with ‘two compartment’ leaf water concepts.     

Oxygen isotope fractionations across individual leaf carbohydrates in grass and tree species

Almost no δ¹⁸O data are available for leaf carbohydrates, leaving a gap in the understanding of the δ¹⁸O relationship between leaf water and cellulose. We measured δ¹⁸O values of bulk leaf water (δ¹⁸O_LW) and individual leaf carbohydrates (e.g. fructose, glucose and sucrose) in grass and tree species and δ¹⁸O of leaf cellulose in grasses. The grasses were grown under two relative humidity (rH) conditions. Sucrose was generally ¹⁸O‐enriched compared with hexoses across all species with an apparent biosynthetic fractionation factor (ε_bio) of more than 27‰ relative to δ¹⁸O_LW, which might be explained by isotopic leaf water and sucrose synthesis gradients. δ¹⁸O_LW and δ¹⁸O values of carbohydrates and cellulose in grasses were strongly related, indicating that the leaf water signal in carbohydrates was transferred to cellulose (ε_bio = 25.1‰). Interestingly, damping factor p_exp_x, which reflects oxygen isotope exchange with less enriched water during cellulose synthesis, responded to rH conditions if modelled from δ¹⁸O_LW but not if modelled directly from δ¹⁸O of individual carbohydrates. We conclude that δ¹⁸O_LW is not always a good substitute for δ¹⁸O of synthesis water due to isotopic leaf water gradients. Thus, compound‐specific δ¹⁸O analyses of individual carbohydrates are helpful to better constrain (post‐)photosynthetic isotope fractionation processes in plants.     

The 18O-signal transfer from water vapour to leaf water and assimilates varies among plant species and growth forms

The ¹⁸O signature of atmospheric water vapour (δ¹⁸O_V) is known to be transferred via leaf water to assimilates. It remains, however, unclear how the ¹⁸O-signal transfer differs among plant species and growth forms. We performed a 9-hr greenhouse fog experiment (relative humidity ≥ 98%) with ¹⁸O-depleted water vapour (−106.7‰) on 140 plant species of eight different growth forms during daytime. We quantified the ¹⁸O-signal transfer by calculating the mean residence time of O in leaf water (MRT_LW) and sugars (MRT_Sugars) and related it to leaf traits and physiological drivers. MRT_LW increased with leaf succulence and thickness, varying between 1.4 and 10.8 hr. MRT_Sugars was shorter in C₃ and C₄ plants than in crassulacean acid metabolism (CAM) plants and highly variable among species and growth forms; MRT_Sugars was shortest for grasses and aquatic plants, intermediate for broadleaf trees, shrubs, and herbs, and longest for conifers, epiphytes, and succulents. Sucrose was more sensitive to δ¹⁸O_V variations than other assimilates. Our comprehensive study shows that plant species and growth forms vary strongly in their sensitivity to δ¹⁸O_V variations, which is important for the interpretation of δ¹⁸O values in plant organic material and compounds and thus for the reconstruction of climatic conditions and plant functional responses.     

Surrounding species diversity improves subtropical seedlings’ carbon dynamics

Increasing biodiversity has been linked to higher primary productivity in terrestrial ecosystems. However, the underlying ecophysiological mechanisms remain poorly understood. We investigated the effects of surrounding species richness (monoculture, two‐ and four‐species mixtures) on the ecophysiology of Lithocarpus glaber seedlings in experimental plots in subtropical China. A natural rain event isotopically labelled both the water uptaken by the L. glaber seedlings and the carbon in new photoassimilates through changes of photosynthetic discrimination. We followed the labelled carbon (C) and oxygen (O) in the plant–soil–atmosphere continuum. We measured gas‐exchange variables (C assimilation, transpiration and above‐ and belowground respiration) and δ¹³C in leaf biomass, phloem, soil microbial biomass, leaf‐ and soil‐respired CO₂ as well as δ¹⁸O in leaf and xylem water. The ¹³C signal in phloem and respired CO₂ in L. glaber in monoculture lagged behind those in species mixture, showing a slower transport of new photoassimilates to and through the phloem in monoculture. Furthermore, leaf‐water ¹⁸O enrichment above the xylem water in L. glaber increased after the rain in lower diversity plots suggesting a lower ability to compensate for increased transpiration. Lithocarpus glaber in monoculture showed higher C assimilation rate and water‐use efficiency. However, these increased C resources did not translate in higher growth of L. glaber in monoculture suggesting the existence of larger nongrowth‐related C sinks in monoculture. These ecophysiological responses of L. glaber, in agreement with current understanding of phloem transport are consistent with a stronger competition for water resources in monoculture than in species mixtures. Therefore, increasing species diversity in the close vicinity of the studied plants appears to alleviate physiological stress induced by water competition and to counterbalance the negative effects of interspecific competition on assimilation rates for L. glaber by allowing a higher fraction of the C assimilated to be allocated to growth in species mixture than in monoculture.     

Stand structure modulates the long-term vulnerability of Pinus halepensis to climatic drought in a semiarid Mediterranean ecosystem

We investigated whether stand structure modulates the long-term physiological performance and growth of Pinus halepensis Mill. in a semiarid Mediterranean ecosystem. Tree radial growth and carbon and oxygen stable isotope composition of latewood (δ(13)C(LW) and δ(18)O(LW), respectively) from 1967 to 2007 were measured in P. halepensis trees from two sharply contrasting stand types: open woodlands with widely scattered trees versus dense afforested stands. In both stand types, tree radial growth, δ(13)C(LW) and δ(18)O(LW) were strongly correlated with annual rainfall, thus indicating that tree performance in this semiarid environment is largely determined by inter-annual changes in water availability. However, trees in dense afforested stands showed consistently higher δ(18)O(LW) and similar δ(13)C(LW) values compared with those in neighbouring open woodlands, indicating lower stomatal conductance and photosynthesis rates in the former, but little difference in water use efficiency between stand types. Trees in dense afforested stands were more water stressed and showed lower radial growth, overall suggesting greater vulnerability to drought and climate aridification compared with trees in open woodlands. In this semiarid ecosystem, the negative impacts of intense inter-tree competition for water on P. halepensis performance clearly outweigh potential benefits derived from enhanced infiltration and reduced run-off losses in dense afforested stands.     

The “isohydric trap”: A proposed feedback between water shortage,stomatal regulation,and nutrient acquisition drives differential growth and survival of European pines under climatic dryness

Climatic dryness imposes limitations on vascular plant growth by reducing stomatal conductance, thereby decreasing CO₂ uptake and transpiration. Given that transpiration‐driven water flow is required for nutrient uptake, climatic stress‐induced nutrient deficit could be a key mechanism for decreased plant performance under prolonged drought. We propose the existence of an “isohydric trap,” a dryness‐induced detrimental feedback leading to nutrient deficit and stoichiometry imbalance in strict isohydric species. We tested this framework in a common garden experiment with 840 individuals of four ecologically contrasting European pines (Pinus halepensis, P. nigra, P. sylvestris, and P. uncinata) at a site with high temperature and low soil water availability. We measured growth, survival, photochemical efficiency, stem water potentials, leaf isotopic composition (δ¹³C, δ¹⁸O), and nutrient concentrations (C, N, P, K, Zn, Cu). After 2 years, the Mediterranean species Pinus halepensis showed lower δ¹⁸O and higher δ¹³C values than the other species, indicating higher time‐integrated transpiration and water‐use efficiency (WUE), along with lower predawn and midday water potentials, higher photochemical efficiency, higher leaf P, and K concentrations, more balanced N:P and N:K ratios, and much greater dry‐biomass (up to 63‐fold) and survival (100%). Conversely, the more mesic mountain pine species showed higher leaf δ¹⁸O and lower δ¹³C, indicating lower transpiration and WUE, higher water potentials, severe P and K deficiencies and N:P and N:K imbalances, and poorer photochemical efficiency, growth, and survival. These results support our hypothesis that vascular plant species with tight stomatal regulation of transpiration can become trapped in a feedback cycle of nutrient deficit and imbalance that exacerbates the detrimental impacts of climatic dryness on performance. This overlooked feedback mechanism may hamper the ability of isohydric species to respond to ongoing global change, by aggravating the interactive impacts of stoichiometric imbalance and water stress caused by anthropogenic N deposition and hotter droughts, respectively.     

Recent and historical discharge of a large European river system - oxygen isotopic composition of river water and skeletal aragonite of Unionidae in the Rhine

Seasonal variations in the oxygen isotopic composition of Rhine River water were analyzed in detail and compared with the oxygen isotopic record from recent and historical specimens of freshwater bivalves (Unionidae). The purpose of this study was to investigate the potential use of these aragonitic bivalves as proxy recorders for varying amounts and sources of discharge, and thereby infer climate change. Seasonal variations in the river water δ¹⁸O are on the order of 1-2‰. During summer, Alpine melt-water contributes significantly to the total discharge, resulting in average values of −10 to −10.5‰, whereas the non-Alpine contribution is higher during winter, as indicated by mean δ¹⁸O values of −8.5 to −9‰. The basic pattern of the modern seasonal variation of δ¹⁸O of river water can be described by a numerical mass balance approximation of the various contributions from the Alpine and non-Alpine catchments with their average δ¹⁸O composition. The δ¹⁸O of growth increments in the prismatic shell layer of Anodonta corresponds perfectly to what is predicted by known fractionation of ¹⁸O between water and aragonite. Shell growth is restricted to water temperatures above 8-10°C, so variations in δ¹⁸O and the river water temperature are faithfully recorded by relatively large growth increments during summer. The distinctive isotopic signatures of individual flood events during summer and autumn are also recorded in the shells.     

Correlating δ13C and δ18O in cellulose of trees

We measured the carbon and oxygen isotopic composition of stem cellulose of Pinus sylvestris, Picea abies, Fagus sylvatica and Fraxinus excelsior. Several sites along a transect of a small valley in Switzerland were selected which differ in soil moisture conditions. At every site, six trees per species were sampled, and a sample representing a mean value for the period from 1940 to 1990 was analysed. For all species, the mean site δ¹³C and δ¹⁸O of stem cellulose are related to the soil moisture availability, whereby higher isotope ratios are found at drier sites. This result is consistent with isotope fractionation models when assuming enhanced stomatal resistance (thus higher δ¹³C of incorporated carbon) and increased oxygen isotope enrichment in the leaf water (thus higher δ¹⁸O) at the dry sites. δ¹⁸ O-δ¹³C plots reveal a linear relationship between the carbon and oxygen isotopes in cellulose. To interpret this relationship we developed an equation which combines the above-mentioned fractionation models. An important new parameter is the degree to which the leaf water enrichment is reflected in the stem cellulose. In the combined model the slope of the δ¹⁸O-δ¹³C plot is related to the sensitivity of the p_i/p_a of a plant to changing relative humidity.     

Overstorey and juvenile response to thinning and drought in a jarrah (Eucalyptus marginata Donn ex Sm.) forest of southwestern Australia

Aims

Forest thinning is expected to affect tree water use and carbon assimilation, but the related influence from climate variability is little known. Recent forest thinning in the Wungong catchment coincided with a record dry year following the thinning, which provides a rare opportunity to understand the climate influence on the thinning effect.

Methods

A field experiment was conducted to examine changes before and after thinning, especially the rainfall, soil moisture, leaf water status, tissue isotope signature (¹³?C and ¹⁵?N) and N concentration of overstorey and understorey juvenile trees of Eucalyptus marginata (Donn ex Sm.).

Results

Despite the post-thinning drought, surface soil was moister and juvenile jarrah plants were less water stressed, attributable to reduced rain interception and transpiration as a result of less canopy cover. The overstorey was under stress but mainly due to drought rather than by thinning. The concentration of N declined in both tree stems and juvenile leaves along with available N in soil, suggesting a soil N limitation. No treatment effects were detected from leaf relative water content and tissue isotope signature (¹³?C and ¹⁵?N).

Conclusions

The drought effects were superimposed over the thinning effects on overstorey growth, with stemwood δ¹³C being a major indicator of water stress. The water relations and carbon assimilation of understorey juveniles were however dependent more on topsoil moisture, and the wetter soil during the year following thinning enhanced growth activity and hence the depletion of ¹³?C (more negative δ¹³C) in juvenile leaves.     

Temperature versus species-specific influences on the stable oxygen isotope ratio of tree rings

Stable isotopic ratios integrate ecosystem variability while reflecting change in both environmental and biological processes. At sites, where climate does not strongly limit tree growth, co-occurring trees may display large discrepancies in stable oxygen isotopic ratios (δ¹⁸O) due to the interplay between biological processes (competition for light and nutrients, individual tree physiology, etc.) and climate. For a better quantification of the isotope variability within and among trees, the climatic and/or individual tree effects on seasonal δ¹⁸O variations in precipitation, soil water, leaf water and leaf organic material (whole leaf, cellulose and starch) and annual δ¹⁸O variations in tree-ring cellulose for Fagus sylvatica (Fs), Quercus robur (Qr), Carpinus betulus (Cb) and Pinus sylvestris (Ps) were studied in a mature temperate forest in Switzerland, using a mixed linear regression model technique. Furthermore, the influence of environmental factors on δ¹⁸O was assessed by means of three common isotope fractionation models. Our statistical analysis showed that except for Ps, a greater portion of δ¹⁸O variance in leaf compounds can be explained by individual tree effects, compared to temperature. Concerning tree-ring cellulose, only Fs and Ps show a significant temperature signal (maximum 12% of the variance explained), while the individual tree effect significantly explains δ¹⁸O for all species for a period of 38 years. Large species differences resulted in a limited ability of the isotope fractionation models to predict measured values. Overall, we conclude that in a diverse mixed forest stand, individual tree responses reduce the potential extraction of a temperature signal from δ¹⁸O.     

Climatic responses of tree-ring width and δ13C signatures of sessile oak (Quercus petraea Liebl.) on soils with contrasting water supply

We investigated climate–growth relationships (in terms of tree-ring width, basal area increment (BAI), and tree-ring δ¹³C signatures) of Quercus petraea in Central Europe (Luxembourg). Tree responses were assessed for 160 years and compared for sites with contrasting water supply (i.e. Cambisols vs. Regosols with 175 and 42 mm available water capacity, respectively). Oak trees displayed very low climate sensitivity, and climatic variables explained only 24 and 21 % of variance in tree-ring width (TRW) (Cambisol and Regosol sites, respectively). Contrary to our expectations, site-related differences in growth responses (i.e. BAI, δ¹³C signatures) to climate shifts were not significant. This finding suggests a high plasticity of oak trees in the study area. Despite a distinct growth depression found for all trees in the decade 1988–1997 (attributable to increasing annual mean temperatures by 1.1 °C), oak trees completely recovered in subsequent years. This indicates a high resilience of sessile oak to climate change. Shifts in δ¹³C_corr signatures were mainly affected by temperature, and peaks in δ¹³C_corr values (corrected for the anthropogenic increase in atmospheric CO₂) coincided with decadal maximum temperatures. Correlations between δ¹³C signatures and TRW (mainly affected by precipitation) were not significant. This finding suggests that wood growth often was disconnected from carbon assimilation (e.g. due to carbon storage in the trunk or allocation to seeds). Since the selection of drought-resistant tree species gains importance within the context of adaptive forest management strategies, Q. petraea proves to be an adaptive tree species in Central Europe’s forests under shifting climatic conditions.     

Non-climatic variations in the oxygen isotopic compositions of plants

The ¹⁸O content of leaf water strongly influences the ¹⁸O contents of atmospheric CO₂ and O₂. The ¹⁸O signatures of these atmospheric gases, in turn, emerge as important indicators of large-scale gas exchange processes. Better understanding of the factors that influence the isotopic composition of leaf water is still required, however, for the quantitative utilization of these tracers. The ¹⁸O enrichment of leaf water relative to local meteoric water, is known to reflect climatic conditions. Less is known about the extent variations in the ¹⁸O content of leaf water are influenced by nonclimatic, species-specific characteristics. In a collection of 90 plant species from all continents grown under the same climatic conditions in the Jerusalem Botanical Garden we observed variations of about 9‰ in the δ¹⁸O values of stem water, δ_s, and of about 14‰ in the mid-day δ¹⁸O enrichment of bulk leaf water, δ_LW–δ_s. Differences between δ¹⁸O values predicted by a conventional evaporation model, δ_M, and δ_LW ranged between – 3.3‰ and + 11.8‰. The δ¹⁸O values of water in the chloroplasts (δ_ch) in leaves of 10 selected plants were estimated from on-line CO₂ discrimination measurements. Although much uncertainty is still involved in these estimates, the results indicated that δ_ch can significantly deviate from δ_M in species with high leaf peclet number. The δ¹⁸O values of bulk leaf water significantly correlated with δ¹⁸O values of leaf cellulose (directly) and with instantaneous water use efficiency (A/E, inversely). Differences in isotopic characteristics among conventionally defined vegetation types were not significant, except for conifers that significantly differed from shrubs in δ¹⁸O and δ¹³C values of cellulose and in their peclet numbers, and from deciduous woodland species in their δ¹⁸O and δ¹³C values of cellulose. The results indicated that predictions of the δ¹⁸O values of leaf water (δ_LW, δ_M and δ_ch) could be improved by considering plant species-specific characteristics.