Mediterranean evergreen oaks have to survive a long summer drought. Roots may play a relevant role under these conditions. We studied their structure and function in a mature Quercus suber L. tree in central Portugal. The root system was mapped till the lowest water table level (4.5 m depth). Xylem anatomy was analyzed in a vertical profile belowground. Sap flow was continuously monitored for 1.5 yrs in the stem and roots of this intensively studied tree (heat field deformation method) and in the stem of four trees (Granier method), in relation to environmental variables and predawn leaf water potential. The sources of water uptake were assessed by stable isotope analyses in summer. Results showed a dimorphic root system with a network of superficial roots linked to sinker roots, and a taproot diverting into tangles of deep fine roots submerged for long periods, with parenchyma aerenchyma. Transpiration was not restricted in summer due to root access to groundwater. The isotopic d 18 O signature of twig xylem water was similar to that of groundwater in the dry season. Two functional types of superficial roots were identified: shallow connected and deep connected roots. A modeling approach was built considering that each superficial root was linked to a sinker, with part of the root deep connected (between the stem and the sinker) and part shallow connected (between the sinker and topsoil). This conceptual framework simulated tree stem sap flow from root sap flow with a high efficiency (R 2 = 0.85) in four plot trees. On an annual basis, soil water and groundwater contributions were 69.5% and 30.5% of stem flow, respectively. Annual hydraulic lift and hydraulic descent were 0.9% and 37.0% of stem flow, respectively. The trees maximize the exploitation of the environmental resources by using the topsoil water during most of the year, and groundwater together with hydraulic lift (nutrient supply) in the dry summer. This study shows that a dimorphic root system, with roots reaching groundwater, is an efficient strategy of Q. suber trees to cope with seasonal drought. Knowledge of the functional behavior of Q. suber trees under shallow water table conditions may contribute to the definition of better adapted management practices and to anticipate their responses to climate change.

Mediterranean evergreen oak woodlands of southern Portugal (montados) are savannah-type ecosystems with a widely sparse tree cover, over extensive grassland. Therefore, ecosystem water fluxes derive from two quite differentiated sources: the trees and the pasture. Partitioning of fluxes according to these different sources is necessary to quantify overall ecosystem water losses as well as to improve knowledge on its functional behaviour. In southern Iberia, these woodlands are subjected to recurrent droughts. Therefore, reaction/resilience to water stress becomes an essential feature of vegetation on these ecosystems. Long-term tree transpiration was recorded for 6 years from a sample of holm oak (Quercus ilex ssp. rotundifolia) trees, using the Granier sap flow method. Ecosystem transpiration was measured by the eddy covariance technique for an 11-month period (February to December 2005), partly coincident with a drought year. Pasture transpiration was estimated as the difference between ecosystem (eddy covariance) and tree (sap flow) transpiration. Pasture transpiration stopped during the summer, when the surface soil dried up. In the other seasons, pasture transpiration showed a strong dependence on rainfall occurrence and on top soil water. Conversely, trees were able to maintain transpiration throughout the summer due to the deep root access to groundwater. Q. ilex trees showed a high resilience to both seasonal and annual drought. Tree transpiration represented more than half of ecosystem transpiration, in spite of the low tree density (30 trees ha À1) and crown cover fraction (21%). Tree evapotranspiration was dominated by transpiration (76%), and interception loss represented only 24% of overall tree evaporation.

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Quercus suber and Quercus ilex trees are major elements of Mediterranean landscapes, which are threatened by the increasing water deficits related to climate change. To contribute to the understanding of the capacity of these oaks to withstand severe drought we assessed the vulnerability to xylem embolism and the anatomical traits in current-year shoots. Data were collected in mature trees at two sites, in central/coastal and southern/inland Portugal. In situ safety margins to hydraulic failure were evaluated from long-term predawn and midday leaf water potential records. Results showed that xylem vulnerability to embolism was similar in Q. ilex and Q. suber. The 50% loss in hydraulic conductivity (Ψxyl,50PLC) was observed at xylem water potentials of −2.9 and −3.2 MPa in shoots of Q. suber and Q. ilex, respectively. Values of mean vessel diameter of Q. suber shoots at both sites suggest an intra-species adaptation to the local water availability, with larger vessels at the more mesic site. In situ hydraulic safety margins observed in shoots showed that, even during the driest periods, both oaks lived comfortably above the most critical embolism thresholds. However, the hydraulic safety margins were narrower in the driest site. Results are relevant to the understanding of survival, growth, and functional behaviour of evergreen oaks in Mediterranean climates, under recurrent/seasonal drought conditions

Abstract:Water is one of the major environmental factors limiting plant growth and survival in the Mediterranean region. Quercus suber L. woodlands occupy vast areas in the Iberian Peninsula, frequently under shallow water table conditions. The relative magnitude of soil and groundwater uptake to supply transpiration is not easy to evaluate under these circumstances. We recently developed a conceptual framework for the functioning of the root system in Q. suber that simulates well tree transpiration, based on two types of root behaviour: shallow connected and deep connected. Although this significantly improved knowledge on the functional traits of Mediterranean Q. suber, the approach has the limitation of requiring root sap flow data, which are seldom available. In this work, we present alternative methodologies to assess if trees are connected to groundwater and to estimate the soil and groundwater contributions to tree transpiration. We provide evidence on the tree unrestricted access to groundwater solely based on meteorological, stem sap flow and leaf water potential data. Using a soil mass balance approach, we estimated the yearly soil and groundwater contributions to tree transpiration: 69.7% and 30.3%, respectively. Groundwater uptake became dominant in the dry summer: 73.2% of tree transpiration. Results reproduce extremely well those derived from root modelling. Because of its simplicity both in formulation and data requirements, our approach is potentially liable to be adapted to other groundwaterdependent Mediterranean oak sites, where interactions between land use and water resources may be relevant.

The topology of the xylem network is likely to affect the transport of water, propagation of embolism and plant survival and growth. Few studies have been conducted on the hydraulics of the entire water pathway in trees. We evaluated the hydraulic connections from roots to branches in a mature Quercus suber L. tree, through sap flow responses upon branch severing. Sap flow was recorded in branches, stem and roots by the heat field deformation (HFD) method. Results showed that roots, except for the taproot, were hydraulically connected to all branches, but the rest of the tree (stem, branches and taproot) was highly sectored. In the large roots that showed an integrated response to branch severing, the outer xylem was preferentially connected to the same side branch and the inner xylem to the opposite branch. The hydraulic sectoriality in branches, stem and taproot may be regarded as an adaptive trait to water stress. The integrated hydraulic structure of roots is advantageous under patchy soil conditions, but may allow the spread of root diseases. The HFD sap flow method proved extremely useful to calculate xylem flux connectivity between different organs of a large tree, providing a comprehensive picture of its hydraulic architecture.

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Aim of the study: Water scarcity is the main limitation to forest growth and tree survival in the Mediterranean hot climate zone. This paper reviews literature on the relations between water and forests in the region, and their implications on forest and water resources management. The analysis is based on a hydraulic interpretation of tree functioning.Area of the study: The review covers research carried out in the Mediterranean hot climate zone, put into perspective of wider/ global research on the subject. The scales of analysis range from the tree to catchment levels.Material and methods: For literature review we used Scopus, Web of Science and Google Scholar as bibliographic databases. Data from two Quercus suber sites in Portugal were used for illustrative purposes.Main results: We identify knowledge gaps and discuss options to better adapt forest management to climate change under a tree water use/availability perspective. Forest management is also discussed within the wider context of catchment water balance: water is a constraint for biomass production, but also for other human activities such as urban supply, industry and irrigated agriculture.Research highlights: Given the scarce and variable (in space and in time) water availability in the region, further research is needed on: mapping the spatial heterogeneity of water availability to trees; adjustment of tree density to local conditions; silvicultural practices that do not damage soil properties or roots; irrigation of forest plantations in some specific areas; tree breeding. Also, a closer cooperation between forest and water managers is needed.