Vegetation in water‐limited ecosystems relies strongly on access to deep water reserves to withstand dry periods. Most of these ecosystems have shallow soils over deep groundwater reserves. Understanding the functioning and functional plasticity of species‐specific root systems and the patterns of or differences in the use of water sources under more frequent or intense droughts is therefore necessary to properly predict the responses of seasonally dry ecosystems to future climate. We used stable isotopes to investigate the seasonal patterns of water uptake by a sclerophyll forest on sloped terrain with shallow soils. We assessed the effect of a long‐term experimental drought (12 years) and the added impact of an extreme natural drought that produced widespread tree mortality and crown defoliation. The dominant species, Quercus ilex, Arbutus unedo and Phillyrea latifolia, all have dimorphic root systems enabling them to access different water sources in space and time. The plants extracted water mainly from the soil in the cold and wet seasons but increased their use of groundwater during the summer drought. Interestingly, the plants subjected to the long‐term experimental drought shifted water uptake toward deeper (10–35 cm) soil layers during the wet season and reduced groundwater uptake in summer, indicating plasticity in the functional distribution of fine roots that dampened the effect of our experimental drought over the long term. An extreme drought in 2011, however, further reduced the contribution of deep soil layers and groundwater to transpiration, which resulted in greater crown defoliation in the drought‐affected plants. This study suggests that extreme droughts aggravate moderate but persistent drier conditions (simulated by our manipulation) and may lead to the depletion of water from groundwater reservoirs and weathered bedrock, threatening the preservation of these Mediterranean ecosystems in their current structures and compositions.
Summary Stable isotopes are extensively used as tracers for the study of plant‐water sources. Isotope‐ratio infrared spectroscopy (IRIS) offers a cheaper alternative to isotope‐ratio mass spectroscopy (IRMS), but its use in studying plant and soil water is limited by the spectral interference caused by organic contaminants. Here, we examine two approaches to cope with contaminated samples in IRIS: on‐line oxidation of organic compounds (MCM) and post‐processing correction. We assessed these methods compared to IRMS across 136 samples of xylem and soil water, and a set of ethanol– and methanol–water mixtures. A post‐processing correction significantly improved IRIS accuracy in both natural samples and alcohol dilutions, being effective with concentrations up to 8% of ethanol and 0.4% of methanol. MCM outperformed the post‐processing correction in removing methanol interference, but did not effectively remove interference for high concentrations of ethanol. By using both approaches, IRIS can overcome with reasonable accuracy the analytical uncertainties associated with most organic contaminants found in soil and xylem water. We recommend the post‐processing correction as the first choice for analysis of samples of unknown contamination. Nevertheless, MCM can be more effective for evaluating samples containing contaminants responsible for strong spectral interferences at low concentrations, such as methanol.
Forests play a key role in the carbon balance of terrestrial ecosystems. One of the main uncertainties in global change predictions lies in how the spatiotemporal dynamics of forest productivity will be affected by climate warming. Here we show an increasing influence of climate on the spatial variability of tree growth during the last 120 y, ultimately leading to unprecedented temporal coherence in ring-width records over wide geographical scales (spatial synchrony). Synchrony in growth patterns across cold-constrained (central Siberia) and droughtconstrained (Spain) Eurasian conifer forests have peaked in the early 21st century at subcontinental scales (∼1,000 km). Such enhanced synchrony is similar to that observed in trees co-occurring within a stand. In boreal forests, the combined effects of recent warming and increasing intensity of climate extremes are enhancing synchrony through an earlier start of wood formation and a stronger impact of year-to-year fluctuations of growing-season temperatures on growth. In Mediterranean forests, the impact of warming on synchrony is related mainly to an advanced onset of growth and the strengthening of droughtinduced growth limitations. Spatial patterns of enhanced synchrony represent early warning signals of climate change impacts on forest ecosystems at subcontinental scales. U nderstanding how climate change affects forests across multiple spatiotemporal scales is important for anticipating its impacts on terrestrial ecosystems. Increases in atmospheric CO 2 concentration and shifts in phenology (1-3) could favor tree growth by enhancing photosynthesis and extending the effective growing period, respectively (4). Conversely, recent warming could increase respiration rates and, together with increasing heat and drought stresses, exert negative impacts on forest productivity (5, 6). Given the uncertainty as to what extent enhanced carbon uptake could be offset by the detrimental effects of warming on tree performance, the actual consequences of climate change on forest carbon cycling remain under debate. Notably, climate change has a stronger impact on forests constrained by climatic stressors, such as suboptimal temperatures or water shortage (7). As high-resolution repositories of biological responses to the environment, dendrochronological archives can be used to monitor this impact (8).The concept of spatial synchrony in tree growth refers to the extent of coincident changes in ring-width patterns among geographically disjunct tree populations (9). Climatic restrictions tend to strengthen growth-climate relationships, resulting in enhanced common ringwidth signals (i.e., more synchronous tree growth). Thus, regional bioclimatic patterns can be delineated by identifying groups of trees whose growth is synchronously driven by certain climatic constraints (10, 11). Previous synthesis studies have provided evidence for globally coherent multispecies responses to climate change in natural systems, including forests, with a focus on the role of increasingly warmer tempe...
Holm oak (Quercus ilexL.) and Aleppo pine (Pinus halepensisMill) are representative of two different functional types of trees extensively found in the Mediterranean: evergreen sclerophyllous and drought-adapted conifers. The former is considered a partially drought-tolerant species, whereas the latter is a typically drought-avoiding, water-saving species. We postulated that contrasting strategies in response to water deficits in Q. ilex and P. halepensis would lead to a differential sensitivity to changes in water availability. To test this hypothesis, we compared the response of both species in growth rate (measured as radial increments) and intrinsic water use efficiency WUE(i), as inferred from carbon isotope discrimination (Delta(13)C) in wood samples] among sites from different provenance regions in NE Spain. We found significant differences in Delta(13)C and growth among provenance regions, partly explained by contrasting water availability. Wood Delta(13)C was positively related with precipitation and the ratio between precipitation and potential evapotranspiration (P/ E). However, these relationships were stronger in P. halepensis (for P / E, r(2)=0.67, P <0.001) than in Q. ilex ( r(2)=0.42, P <0.01). In addition, radial growth was positively related with precipitation and Delta(13)C in P. halepensis ( r(2)=0.32 and r(2)=0.35, respectively, P <0.01), but not in Q. ilex. We concluded that P. halepensis was more sensitive than Q. ilex to water availability, showing faster increase in WUE(i) in response to water stress. We also found that the effect of north/south aspect on Delta(13)C and growth was site-specific, and unrelated to climatic variables.
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