Argania spinosa (the argan tree) is a slowgrowing tree endemic of Morocco, growing on semiarid areas where no other tree species can live. With the aim of predicting temporal changes in A. spinosa woodlands under a probable increase in aridity, we set off to investigate these questions: how do A. spinosa physiological attributes respond to variations in climatic conditions and seasonality, and which is the set of attributes that most affects tree response to environmental conditions? In three study sites, Beni Snassen (North), High-Atlas (Mountain) and Admine Forest in Agadir (Coastal), gas exchange measurements, photochemical efficiency, leaf water potential and different leaf attributes were monitored in February, July and November of 2006. The Mountain site presents the most continental climate. Trees in this site were the most stressed in summer, having the lowest midday leaf water potential values, photochemical efficiency and assimilation rates. We found a Ψ md threshold around -4 MPa, below which stomatal conductance responds linearly to Ψ md . Plants from the North area never reached this threshold during the study period. Although leaf pigments presented a clear seasonal pattern, leaves from Coastal trees exhibit the highest content for each season. The three study sites were separated by two discriminate functions obtained by canonical discriminant analysis. In summer, the Mountain population is separated from the other sites mainly by assimilation rate and F v /F m , while in winter transpiration rates and chlorophyll content are the main discriminant variables. Our study shows that A. spinosa trees adjust their physiological status and leaf attributes to environmental conditions allowing plants to thrive under a dry climate. Under a scenario of global change, the distribution of the argan tree likely shifts to milder areas.

Abstract. This study attempts to identify Plant Functional Types (PFTs) within the scrub vegetation of the stabilized sand dunes of the Doñana National Park (southwestern Spain) and to explore possible links with ecosystem function. Two 10‐km long transects were sampled along a gradient of water table depth from elevated and dry dunes to a border area close to the salt marsh. A matrix of cover values for 20 woody species × 58 plots and a matrix of 24 morphological and physiological traits × 20 species were analysed by means of DCA and TWINSPAN to identify the main vegetation types and PFTs. In order to know the predominant plant traits, the matrix of 20 species × 58 plots was multiplied by the 24 traits × 20 species matrix. The resulting 24 traits × 58 plots matrix was analysed by means of DCA. The main vegetation types coincide with the previous descriptions of the Park with Juniperus phoenicea woodland and xerophytic species dominating the top of the dunes and hygrophytic species with isolated Quercus suber trees in the depressions. Previous classifications of Mediterranean woody plants resulted in two main PFTs. The present analysis confirmed these two groups: maquis versus garrigue, but added four new groups. The analysis of the traits × plots matrix revealed that the main trend of variation is related to the water table gradient, while the traits related to the species of mature plant communities had little weight in the analysis because these communities are restricted to isolated patches.

The Dune System of Don˜ana National Park (SW Spain) exhibit a mosaic of environmental characteristics, with different plant communities, all under the same Mediterranean climate, creating an interesting field laboratory for the study of plant responses to stressing conditions. Fourteen woody plant populations were selected, belonging to either xerophytic or hygrophytic plant communities on stabilised dunes, where topography causes differences in soil water availability. Plants were tagged and morphological and ecophysiological measurements were recorded in winter and summer of 1999. Seasonal differences in ecophysiological measurements together with morphological variables were used as plant traits to identify the main adaptive responses of the species. Cluster analysis of traits separated three groups of plant strategies: spiny legume species; sclerophyll, and semideciduous species. In addition, another two kind of strategies have been found in the semideciduous group of species, those withstanding water shortages, attaining very negative water potentials, low photochemical efficiency, and leaf proline accumulation in summer, as opposed to those tolerating water deficit, with moderate seasonal differences in water potential, proline content and photochemical efficiency. The results of this study indicate that legume species behave as a different functional group and drought-semideciduous species present different adaptive responses under the same environmental stress. Ecophysiological measurements must be used as plant traits to detect functional groups under Mediterranean climate.

Under natural conditions the overlapping of multiple stressors may initiate coordinated ecophysiological responses in Mediterranean species. Seasonal plasticity may enable plants to better cope with adverse environmental conditions and/or resource variability. In this article, we study the seasonal responses of 12 woody species in two sites of differing water availability, in a Mediterranean-type climate. Plants were measured for water potential, photochemical efficiency, photosynthetic pigments and leaf proline content throughout the year. The results revealed that species presented different ecophysiological strategies, even when sharing the same area. In the xerophytic site, some species suffered severe water stress (-12 MPa and F v /F m lower than 0.3), while others exhibited optimal values of F v /F m and only moderate water stress. All the plants recovered after the first autumn rains. In the hygrophytic site, some sclerophyll species did not exhibit signs of water stress, but did suffer photoinhibition in clear winter days. A plasticity index was calculated to provide an integrated value of species plasticity. In summer, plasticity was higher in the xerophytic site, while in winter it was higher in the hygrophytic site. Ordination analysis of the physiological traits supports the traditional gradient of Mediterranean strategies from drought semideciduous to evergreen sclerophyll species, although spiny legume species formed an independent functional group. The functional responses of species clearly differ among plant communities according to the prevailing site stressors, but no unique pattern emerges. Species combine traits in broader strategies according to previous evolutionary story exhibiting a certain amount of trade among traits, each contributing to alleviate a part of the plant stress.

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In South-western Morocco, water scarcity and high temperature are the main factors determining species survival. Argania spinosa (L.) Skeels is a tree species, endemic to Morocco, which is suffering from ongoing habitat shrinkage. Argan trees play essential local ecological and economic roles: protecting soils from erosion, shading different types of crops, helping maintain soil fertility and, even more importantly, its seeds are used by the local population for oil production, with valuable nutritional, medicinal and cosmetic purposes. The main objective of this study was to identify the sources of water used by this species and to assess the effect of water availability on the photosynthetic rate and stem water potential in two populations: one growing on the coast and a second one 10 km inland. Stem water potential, photosynthetic rate and xylem water isotopic composition (δ18O) were seasonally monitored during 2 years. Trees from both populations showed a similar strategy in the use of the available water sources, which was strongly dependent on deep soil water throughout the year. Nevertheless, during the wet season or under low precipitation a more complex water uptake pattern was found with a mixture of water sources, including precipitation and soil at different depths. No evidence was found of the use of either groundwater or atmospheric water in this species. Despite the similar water-use strategy, the results indicate that Argania trees from the inland population explored deeper layers than coastal ones as suggested by more depleted δ18O values recorded in the inland trees and better photosynthetic performance, hence suggesting that the coastal population of A. spinosa could be subjected to higher stress.