<?xml version="1.0" encoding="utf-8" standalone="no"?>
<!DOCTYPE article SYSTEM "http://www.biogeosciences.net/inc/bg/copernicus.dtd">
<article language="en">
	<journal>
		<journal_title>Biogeosciences</journal_title>
		<journal_url>www.biogeosciences.net</journal_url>
		<issn>1726-4170</issn>
		<eissn>1726-4189</eissn>
		<volume_number>6</volume_number>
		<issue_number>11</issue_number>
		<publication_year>2009</publication_year>
	</journal>
	<doi>10.5194/bg-6-2599-2009</doi>
	<article_url>http://www.biogeosciences.net/6/2599/2009/</article_url>
	<abstract_html>http://www.biogeosciences.net/6/2599/2009/bg-6-2599-2009.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/6/2599/2009/bg-6-2599-2009.pdf</fulltext_pdf>
	<start_page>2599</start_page>
	<end_page>2610</end_page>
	<publication_date>2009-11-17</publication_date>
	<article_title content_type="html">An integrated approach shows different use of water resources from Mediterranean maquis species in a coastal dune ecosystem</article_title>
	<authors>
		<author numeration="1" affiliations="1,3">
			<name>S. Mereu</name>
			<email>simone.mereu@libero.it</email>
		</author>
		<author numeration="2" affiliations="1">
			<name>E. Salvatori</name>
		</author>
		<author numeration="3" affiliations="1">
			<name>L. Fusaro</name>
		</author>
		<author numeration="4" affiliations="2">
			<name>G. Gerosa</name>
		</author>
		<author numeration="5" affiliations="3">
			<name>B. Muys</name>
		</author>
		<author numeration="6" affiliations="1">
			<name>F. Manes</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Sapienza University of Rome, Department of Plant Biology, P.le Aldo Moro 5, 00185, Rome, Italy</affiliation>
		<affiliation numeration="2" content_type="html">Catholic University of the Sacred Heart, Department of Mathematics and Physics, Via Musei 41, 25121 Brescia, BS, Italy</affiliation>
		<affiliation numeration="3" content_type="html">Katholieke Universiteit Leuven, Department Forest, Nature and Landscape, Celestijnenlaan 200E, 3001, Leuven, Belgium</affiliation>
	</affiliations>
	<abstract content_type="html">An integrated approach has been used to analyse the dependence of three
Mediterranean species, &lt;i&gt;A. unedo&lt;/i&gt; L., &lt;i&gt;Q. ilex&lt;/i&gt; L., and
&lt;i&gt;P. latifolia&lt;/i&gt; L. co-occurring in a coastal dune ecosystem on two
different water resources: groundwater and rainfed upper soil layers. The
approach included leaf level gas exchanges, sap flow measurements and
structural adaptations between 15 May and 31 July 2007. During this period it
was possible to capture different species-specific response patterns to an
environment characterized by a sandy soil, with a low water retention
capacity, and the presence of a water table. The latter did not completely
prevent the development of a drought response and, combined with previous
studies in the same area, response differences between species have been
partially attributed to different root distributions. Sap flow of
&lt;i&gt;A. unedo&lt;/i&gt; decreased rapidly with the decline of soil water content,
while that of &lt;i&gt;Q. ilex&lt;/i&gt; decreased only moderately. Midday leaf water
potential of &lt;i&gt;P. latifolia&lt;/i&gt; and &lt;i&gt;A. unedo&lt;/i&gt; ranged between
&amp;minus;2.2 and &amp;minus;2.7 MPa throughout the measuring period, while in
&lt;i&gt;Q. ilex&lt;/i&gt; it decreased down to &amp;minus;3.4 MPa at the end of the season.
&lt;i&gt;A. unedo&lt;/i&gt; was the only species that responded to drought with a
decrease of its leaf area to sapwood area ratio from 23.9&amp;plusmn;1.2 (May) to
15.2&amp;plusmn;1.5 (July). While &lt;i&gt;A. unedo&lt;/i&gt; also underwent an almost
stepwise loss on hydraulic conductivity, such a loss did not occur for
&lt;i&gt;Q. ilex&lt;/i&gt;, whereas &lt;i&gt;P. latifolia&lt;/i&gt; was able to slightly increase
its hydraulic conducitivity. These differences show how different plant
compartments coordinate differently between species in their responses to
drought. The different responses appear to be mediated by different root
distributions of the species and their relative resistances to drought are
likely to depend on the duration of the periods in which water remains
extractable in the upper soil layers.</abstract>
	<references>
		<reference numeration="1" content_type="text"> Alley, R. B., Marotzke, J., Nordhaus, W. D., Overpeck, J. T., Peteet, D. M., Pielke Jr., R. A., Pierrehumbert, R. T., Rhines, P. B., Stocker, T. F., Talley, L. D., and Wallace, J. M.: Abrupt climate change, Science, 299, 2005–2010, 2003. </reference>
		<reference numeration="2" content_type="text"> Alessio, G. A., De Lillis, M., Brugnoli, E., and Lauteri, M.: Water sources and water-use efficiency in Mediterranean coastal dune vegetation, Plant Biol., 6, 350–357, 2004. </reference>
		<reference numeration="3" content_type="text"> Asensio, D., Peñuelas, J., Llusia, J., Ogaya, R., and Llusià, J.: Seasonal soil and leaf CO&lt;sub&gt;2&lt;/sub&gt; exchange rates in a Mediterranean holm oak forest and their responses to drought conditions, Atmos. Environ., 41, 2447–2455, 2007. </reference>
		<reference numeration="4" content_type="text"> Bréda, N., Huc, R., Granier, A., and Dreyer, E.: Temperate forest trees and stands under severe drought: a review of ecophysiological responses, adaptation processes and long-term consequences, Ann. For. Sci., 63, 625–644, 2006. </reference>
		<reference numeration="5" content_type="text"> Bucci, S. J., Goldstein, G., Meinzer, F. C., Scholz, F. G., Franco, A. C., and Bustamante, M.: Functional convergence in hydraulic architecture and water relations of tropical savannah trees: from leaf to whole plant, Tree Physiol., 24, 891–899, 2004. </reference>
		<reference numeration="6" content_type="text"> Bussotti, F., Bettini, D., Grossoni, P., Mansuino, S., Nibbi, R., Soda, C., and Tani, C.: Structural and functional traits of \textitQuercus ilex in response to water availability, Environ. Exp. Bot., 47, 11–23, 2002. </reference>
		<reference numeration="7" content_type="text"> Busuoli, G., Bucci, M., and Grillini, M.: Studi geologici, geomorfologici ed idrologici sulla Tenuta Presidenziale di Castelporziano, in: Il sistema ambientale della Tenuta di Castelporziano. Ricerche sulla complessità di un ecosistema forestale costiero Mediterraneo, Rome, Italy, Accademia Nazionale delle Scienze detta dei Quaranta, Scritti e documenti XXVI, Vol 1, 123–157, 2001. </reference>
		<reference numeration="8" content_type="text"> Castillo, J. M., Rubio Casal, A. E., Luque, T., and Figueroa, M. E.: Comparative field summer stress of three tree species co-occurring in Mediterranean coastal dunes, Photosynthetica, 40, 49–56, 2002. </reference>
		<reference numeration="9" content_type="text"> Castro-Díez, P. and Montserrat-Martí, G.: Phenological pattern of fifteen Mediterranean phanaerophytes from \textitQuercus ilex communities of NE-Spain, Plant Ecol., 139, 103–112, 1998. </reference>
		<reference numeration="10" content_type="text"> $\check\rm C$ermák, J., Ku$\check\rm c$era, J., and Nadezhdina, N.: Sap flow measurements with some thermodynamic methods, flow integration within trees and scaling up from sample trees to entire forest stands, Trees-Struct. Funct., 18, 529–546, 2004. </reference>
		<reference numeration="11" content_type="text"> $\check\rm C$ermák, J. and Nadezhdina, N.: Sapwood as the scaling parameter: defining according to xylem water content or radial pattern of sap flow?, Ann. Sci. Forest., 55, 509–521, 1998. </reference>
		<reference numeration="12" content_type="text"> Corcuera, L., Camarero, J. J., and Gil-Pelegrín, E.: Effects of a severe drought on \textitQuercus ilex radial growth and xylem anatomy, Trees, 18, 83–92, 2004. </reference>
		<reference numeration="13" content_type="text"> de Lillis, M.: An ecomorphological study of the evergreen leaf, Braun-Blanquetia, 7, 1–127,1991. </reference>
		<reference numeration="14" content_type="text"> Fares, S., Mereu, S., Scarascia Mugnozza, G., Vitale, M., Manes, F., Frattoni, M., Ciccioli, P., Gerosa, G., and Loreto, F.: The ACCENT-VOCBAS field campaign on biosphere-atmosphere interactions in a Mediterranean ecosystem of Castelporziano (Rome): site characteristics, climatic and meteorological conditions, and eco-physiology of vegetation, Biogeosciences, 6, 1043–1058, 2009. </reference>
		<reference numeration="15" content_type="text"> Ferretti, M., Fagnano, M., Amoriello, T., Badiani, M., Ballarin-Denti, A., Buffoni, A., Bussotti, F., Castagna, A., Cieslik, S., Costantini, A., De Marco, A., Gerosa, G., Lorenzini, G., Manes, F., Merola, G., Nali, C., Paoletti, E., Petriccione, B., Racalbuto, S., Rana, G., Ranieri, A., Tagliaferri, A., Vialetto, G., and Vitale, M.: Measuring, modelling and testing ozone exposure, flux and effects on vegetation in southern European conditions – what does not work. A review from Italy, Environ. Pollut., 146, 648–658, 2007. </reference>
		<reference numeration="16" content_type="text"> Filella, I., Llusià, J., Piñol, J., and Peñuelas, J.: Leaf gas exchange and fluorescence of \textitPhillyrea latifolia, \textitPistacia lentiscus and \textitQuercus ilex saplings in severe drought and high temperature conditions, Environ. Exp. Bot., 39, 213–220, 1998. </reference>
		<reference numeration="17" content_type="text"> Franks, P. J., Drake, P. L., and Froend, R. H.: Anisohydric but isohydrodynamic: seasonally constant plant water potential gradient explained by a stomatal control mechanism incorporating variable plant hydraulic conductance, Plant Cell Environ., 30, 19–30, 2007. </reference>
		<reference numeration="18" content_type="text"> Galmés, J., Pou, A., Alsina, M. M., Tomas, M., Medrano, H., and Flexas, J.: Aquaporin expression in response to different water stress intensities and recovery in Richter-110 \textitVitis sp.: relationship with ecophysiological status, Planta, 226, 671–681, 2007. </reference>
		<reference numeration="19" content_type="text"> Gartner, B. L.: Patterns of xylem variation within a tree and their hydraulic and mechanical consequences. In: Gartner BL ed Plant stems: physiology and functional morphology, Academic, San Diego, 125–145, 1995. </reference>
		<reference numeration="20" content_type="text"> Gratani, L. and Bombelli, A.: Leaf anatomy, inclination, and gas exchange relationships in evergreen schlerophyllous and drought semidiciduos shrub species, Photosynthetica, 374, 573–585, 1999. </reference>
		<reference numeration="21" content_type="text"> Gratani, L. and Varone, L.: Adaptive photosynthetic strategies of the Mediterranean maquis species according to their origin, Photosynthetica, 42, 551–558, 2004. </reference>
		<reference numeration="22" content_type="text"> Jones, H. G.: Plants and microclimate a quantitative approach to environmental plant physiology, Cambridge, Cambridge University Press, 1–384, 1992. </reference>
		<reference numeration="23" content_type="text"> IPCC: Climate Change 2007: Synthesis Report. Contribution of Working Groups I, II and III to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change, edited by: Core Writing Team, Pachauri, R. K., and Reisinger, A., IPCC, Geneva, Switzerland, 104~pp., 2007 </reference>
		<reference numeration="24" content_type="text"> Kaldenhoff, R., Miquel Ribas-Carbo, M., Flexas Sans, J., Lovisolo, C., Heckwolf, M., and Uehlein, N.: Aquaporins and plant water balance, Plant Cell Environ., 31, 658–666, 2008. </reference>
		<reference numeration="25" content_type="text"> Lundblad, M. and Lindroth, A.: Stand transpiration and sapflow density in relation to weather, soil moisture and stand characteristics, Basic Appl. Ecol., 3, 229–243, 2002. </reference>
		<reference numeration="26" content_type="text"> Magnani, F., Grace, J., and Borghetti, M.: Adjustment of tree structure in response to the environment under hydraulic constraints, Funct. Ecol., 16, 385–393, 2002. </reference>
		<reference numeration="27" content_type="text"> Manes, F., Seufert, G., and Vitale, M.: Ecophysiological studies of Mediterranean plant species at the Castelporziano Estate, Atmos. Environ., 31, 51–60, 1997a. </reference>
		<reference numeration="28" content_type="text"> Manes, F., Astorino, G., Vitale, M., and Loreto, F.: Morphofunctional characteristics of \textitQuercus ilex L. leaves of different age and their ecophysiological behaviour during different seasons, Plant. Biosyst., 131, 149–158, 1997c. </reference>
		<reference numeration="29" content_type="text"> Manes, F., Grignetti, A., Tinelli, A., Lenz, R., and Ciccioli, P.: General features of the Castelporziano test site, Atmos. Environ., 31, 19–25, 1997b. </reference>
		<reference numeration="30" content_type="text"> Manes, F., Vitale, M., Donato, E., Giannini, M., and Puppi, G.: Different ability of three Mediterranean oak species to tolerate progressive water stress, Photosynthetica, 44, 387–393, 2006. </reference>
		<reference numeration="31" content_type="text"> Martínez-Vilalta, J., Prat, E., Oliveras, I., and Piñol, J.: Xylem hydraulic properties of roots and stems of nine Mediterranean woody species, Oecologia, 133, 19–29, 2002. </reference>
		<reference numeration="32" content_type="text"> Martínez-Vilalta, J., Mangirón, M., Ogaya, R., Sauret, M., Serrano, L., Peñuelas, J., and Piñol, J.: Sap flow of three co-occurring Mediterranean woody species under varying atmospheric and soil water conditions, Tree Physiol., 23, 747–758, 2003. </reference>
		<reference numeration="33" content_type="text"> Maseda, P. and Fernandez, R.: Stay wet or else: three ways in which plants can adjust to their environment, J. Exp. Bot., 57(15), 3963–3977, 2006. </reference>
		<reference numeration="34" content_type="text"> Mencuccini, M.: The Ecological significance of long-distance water transport: short-term regulation, long-term acclimation and the hydraulic costs of stature across plant life forms, Plant Cell Environ., 26, 163–182, 2003. </reference>
		<reference numeration="35" content_type="text"> Mencuccini, M. and Grace, J.: Climate influences the leaf area/sapwood area ratio in Scots pine, Tree Physiol., 15, 1–10, 1995. </reference>
		<reference numeration="36" content_type="text"> Murray, F. W.: On the computation of saturation vapor pressure, J. Appl. Meteorol., 6, 203–204, 1967. </reference>
		<reference numeration="37" content_type="text"> Nadezhdina, N., Ferreira, M. I., Silva, R., and Pacheco, C. A.: Seasonal variation of water uptake of a \textitQuercus suber tree in Central Portugal, Plant Soil, 305, 105–119, 2008. </reference>
		<reference numeration="38" content_type="text"> Nadezhdina, N., Tributsch, H., and $\check\rm C$ermák, J.: Infra-red images of heat field around a linear heater and sap flow in stems of lime trees under natural and experimental conditions, Ann. For. Sci., 61, 203–213, 2004. </reference>
		<reference numeration="39" content_type="text"> Nicholls, R. J., Wong, P. P., Burkett, V. R., Codignotto, J. O., Hay, J. E., McLean, R. F., Ragoonaden, S., and Woodroffe, C. D.: Coastal systems and low-lying areas, in: Climate Change 2007: Impacts, Adaptation and Vulnerability. Contribution of Working Group II to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change, edited by: Parry, M. L., Canziani, O. F., Palutikof, J. P., van der Linden, P. J., and Hanson, C. E., Cambridge University Press, Cambridge, UK, 315–356, 2007 </reference>
		<reference numeration="40" content_type="text"> Ogaya, R., Peñuelas, J., Martínez-Vilalta, J., and Mangirón, M.: Effects of drought on diameter increment of \textitQuercus ilex, \textitPhillyrea latifolia and \textitArbutus unedo in a holm oak forest of NE Spain, Forest. Ecol. Manag., 180, 175–184, 2003. </reference>
		<reference numeration="41" content_type="text"> Ogaya, R. and Peñuelas, J.: Comparative field study of \textitQuercus ilex and \textitPhillyrea latifolia: photosynthetic response to experimental drought conditions, Environ. Exp. Bot., 50, 137–148, 2003. </reference>
		<reference numeration="42" content_type="text"> Pataki, D. E., Oren, R., Katul, G., and Sigmon, J.: Canopy conductance of \textitPinus taeda, \textitLiquidambar styraciflua and \textitQuercus phellos under varying atmospheric and soil water conditions, Tree Physiol., 18, 307–315, 1998. </reference>
		<reference numeration="43" content_type="text"> Peñuelas, J., Filella, I., Lloret, F., Piñol, J., and Siscart, D.: Effects of a severe drought on water and nitrogen use by \textitQuercus ilex and \textitPhyllirea latifolia, Biol. Plantarum, 43, 47–53, 2000. </reference>
		<reference numeration="44" content_type="text"> Peñuelas, J., Filella, I., Llusia, J., Siscart, D., and Piñol, J.: Comparative field study of spring and summer leaf gas exchange and photobiology of the mediterranean trees \textitQuercus ilex and \textitPhillyrea latifolia, J. Exp. Bot., 49, 229–238, 1998. </reference>
		<reference numeration="45" content_type="text"> Pignatti, S., Bianco, P. M., Tescarollo, P., and Scarascia Mugnozza, G. T.: La vegetazione della tenuta di Castelporziano. In: Il sistema ambientale della tenuta di presidenziale di Castelporziano, Accademia Nazionale dei Quaranta, &quot;scritti e documenti&quot; XXVI, Roma, Vol II, 441–709, 2001. </reference>
		<reference numeration="46" content_type="text"> Pitacco, A., Gallinaro, N., and Giulivo, C.: Evaluation of actual evapotraspiration of a \textitQuercus ilex L. stand by the Bowen ratio Energy budget method, Vegetatio, 99–100, 163–168, 1992. </reference>
		<reference numeration="47" content_type="text"> Sánchez-Vilas, J. and Retuerto, R.: \textitQuercus ilex shows significant among-population variability in functional and growth traits but maintains invariant scaling relations in biomass allocation, Int. J. Plant. Sci., 168, 973–983, 2007. </reference>
		<reference numeration="48" content_type="text"> Schofield, R. K.: The pF of water in the soil, in: Trans. Int. Congr. Soil Sci., Crowther, E. M., et al., Thomas Murby &amp; Co., Oxford, London, UK, 2, 37–48, 30 July–7 August 1935. </reference>
		<reference numeration="49" content_type="text"> Schultz, H. R.: Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown \textitVitis vinifera L. cultivars during drought, Plant Cell Environ., 26, 1393–1405, 2003. </reference>
		<reference numeration="50" content_type="text"> Serrano, L., Peñuelas, J., Ogaya, R., and Savé, R.: Tissue-water relations of two co-occurring evergreen Mediterranean species in response to seasonal and experimental drought conditions, J. Plant. Res., 118, 263–269, 2005. </reference>
		<reference numeration="51" content_type="text"> Serrano, L. and Peñuelas, J.: Contribution of physiological and morphological adjustments to drought resistance in two Mediterranean tree species, Biol. Plant., 49, 551–559, 2005. </reference>
		<reference numeration="52" content_type="text"> Tardieu, F. and Simonneau, T.: Variability among species of stomatal control under fluctuating soil water status and evaporative demand: modelling isohydric and anisohydric behaviours, J. Exp. Bot., 49, 419–432, 1998. </reference>
		<reference numeration="53" content_type="text"> Tognetti, R., Cherubini, P., and Innes, J.: Comparative stemgrowth rates of Mediterranean trees under background and naturally enhanced ambient CO&lt;sub&gt;2&lt;/sub&gt; concentrations, New Phytol., 146, 59–74, 2000a. </reference>
		<reference numeration="54" content_type="text"> Tognetti, R., Minnocci, A., Peñuelas, J., Raschi, A., and Jones, M. B.: Comparative field water relations of three Mediterranean shrub species co-occurring at a natural CO&lt;sub&gt;2&lt;/sub&gt; vent, J. Exp. Bot., 51, 1135–1146, 2000b. </reference>
		<reference numeration="55" content_type="text"> Tognetti, R., Giovannelli, A., Lavini, A., Morelli, G., Fragnito, F., and D&apos;Andria, R.: Assessing environmental controls over conductances through the soil-plant-atmosphere continuum in an experimental olive tree plantation of southern Italy, Agr. Forest Meteorol., 149, 1229–1249, 2009. </reference>
		<reference numeration="56" content_type="text"> Tretiach, M.: Photosynthesis and transpiration of evergreen Mediterranean and deciduous trees in an ecotone during a growing season, Acta Oecol., 143, 341–360, 1993. </reference>
		<reference numeration="57" content_type="text"> Vitale, M. and Manes, F.: Role of changing environmental parameters in leaf gas exchange of \textitArbutus unedo L. assessed by field and laboratory measurements, Photosynthetica, 43, 99–106, 2005. </reference>
		<reference numeration="58" content_type="text"> Whitehead, D. and Jarvis, P. G.: Coniferous forests and plantations, in: Water deficits and plant growth, edited by: Kozlowski, T. T., Academic Press, New York, NY, VI, 49–152, USA. </reference>
	</references>
</article>

