Suites of plant traits in species from different stages of a Mediterranean secondary succession.

The aim of this study was to detect suites of traits related to whole plant and seed morphology, phenology and resource use--including water--in species differing in successional status. Twenty traits were measured on 55 species representative of 5 successional stages in Mediterranean southern France, including eight pertaining to phenology and five to water economy. Suites of traits that changed along succession in agreement with the acquisition/conservation trade-off were completed by continuous changes in phenology. Early successional species had leaves with a high specific leaf area that were produced and lost continuously through the growing season. Late-successional species were taller with long-lived, high delta(13)C leaves produced during short periods, most of them persisting during summer, and produced large seeds requiring a long ripening period. Replacement of species occurred with change in strategies of drought survival: early successional species escaped drought by dying before summer; later herbaceous species maintained favourable water status in relation to leaf shedding during summer; late successional trees with a large body allowing access to a large pool of resources, produced dense leaves that could tolerate desiccation. These changes occurred concomitantly with a shift in CSR strategies, using traits related to resource use, plant size and flowering phenology: ruderal herbs were replaced by more stress-tolerant herbs and shrubs throughout the succession, with competitive trees dominating the latest successional stage. These results suggest that the breadth of functional variability found in natura is not predicted by the CSR framework, and calls for a more integrated view of whole plant functioning.