The effects of water deficit stress and plant water potential (w) on monoterpene and sesquiterpene leaf emissions from Rosmarinus officinalis, Pinus halepensis, Cistus albidus and Quercus coccifera were studied over 11 days of water withholding (from t 1 to t 11 ), after substrates had achieved their field capacity (control pots: t 0 ). Volatile compounds were sampled from the same twig per plant all throughout the study, using a dynamic bag enclosure system. Volatiles, collected in Tenax TA, were studied by means of GC-FID and GC-MS. Monoterpene emissions of water stressed plants (t 1 -t 11 ) were either similar to those of control seedlings (R. officinalis and Q. coccifera)or higher (P. halepensis and C. albidus). By contrast, sesquiterpene emissions were strongly reduced or inhibited after four days of water withholding, particularly for R. officinalis, thus altering terpene emission composition. Despite the positive effect of water stress on leaf monoterpene emissions of P. halepensis and C. albidus, the significant correlation between these emissions and w showed a slow decrease of these emissions over long term water deficit periods. This contrasted with the rapid decline of sesquiterpene emissions of R. officinalis according to lower values of w. These results provide an overall picture of the different responses of monoterpene and sesquiterpene emissions to progressive water loss. They also reveal the utility of using w for estimating some emission rates of some species according to drought conditions.

Summary1. Over a century of agricultural abandonment across the Mediterranean region has favoured the installation of the pioneer expansionist species Aleppo pine (Pinus halepensis Miller). This species synthesizes a wide range of secondary metabolites that are partially released during needle decomposition, and which can thus affect the 'brown food chain'. Litter decomposition is a key process connecting ecosystem structure and function, and involving microbial and faunal components. 2. The goal of this study was to determine the effect of chemical compounds from Aleppo pine needles on the litter decomposition process along a gradient of Mediterranean forest secondary succession. Using in situ litterbags, we compared the dynamics of decomposers, particularly the relative contributions of fungal and mesofauna biomass to litter mass loss (calculations based on the measured decomposer biomass, published fungal growth efficiency and mesofauna feeding rate), against the dynamics of secondary metabolites associated with decomposed needles in three successional stages (early, middle and late, i.e. pinewoods that were aged 10, 30 and over 60 years old). 3. Our first key finding was that fungi accounted for the largest portion of overall litter mass loss (60-79%) and detritivorous mesofauna contributed to 8-12%. In the early stage of succession, fungal biomass after 6 months of decomposition was lower than in middle and late stages, and may be responsible for the delay in litter colonization by mesofauna. We linked this result to a clearly longer residence time for phenolic compounds in young pine forest, leading to an overall slowdown in the decomposition process. 4. Synthesis. Litter phenolic content emerged as a key functional trait for predicting litter decomposition, delaying the colonization of litter by decomposers in Mediterranean forest ecosystems. Another key finding is that the relative contributions of fungi and detritivores to needle mass loss were different between the successional stages. From the food-web perspective, the organic matter available for higher trophic levels thus remains unchanged beyond 30 years after pine colonization.

Recent economic and social changes in north Mediterranean regions have led to an important rural depopulation. Consequently, meadows developed on abandoned agricultural lands (characterized by high species richness) undergo reforestation. These former fields are mainly colonized by Pinus halepensis Miller, which is known to synthetize a wide range of secondary metabolites, among these, some could influence plant succession through allelopathy. The allelopathic potential of P. halepensis, was tested against two target species (Lactuca sativa L. and Linum strictum L.) with aqueous extracts obtained from different organs (root and needle) taking into account the individual age (±10, ±30 and >60 years old). Root and needle extracts affected differently germination and growth of the two target species, the responses varying with concentration of extracts, age and organs tested. The strongest inhibitory effect was observed on the germination and growth of L. strictum, exposed to needle extracts of young P. halepensis (±10 years old), and root extracts of older P. halepensis (>30 years old). These extracts contained several phenolic acids (e.g. 4-hydroxybenzoic acid and p-coumaric acid), which are known as allelochemicals and their concentrations vary with age and organ tested. Hence, P. halepensis could influence secondary succession through the release of potential allelochemicals in the environment by leaf leachates or root exudates.