A series of communities were established in situ to differentiate the effects of species richness, functional richness and functional group identity on invasibility of Mediterranean annual old fields. We monitored the demographic and vegetative parameters of two exotic annuals introduced as seedlings, Conyza bonariensis and C. canadensis. Community species richness and functional composition determined resistance to invasion by Conyza.Conyza bonariensis biomass decreased with increasing species richness. Legumes increased the biomass and consequently the net fecundity of both Conyza, while survival was favoured by Asteraceae. Communities with fewer Asteraceae and grasses increased the reproductive effort of C. bonariensis. A separate glasshouse experiment using the same species mixes revealed that establishment of Conyza decreased with increasing species richness or when grasses were present. Patterns of Conyza performance are interpreted in the light of measurements of ecosystem functional parameters, making it possible to formulate hypotheses about mechanisms limiting community invasibility.

At present, no efficient technique is available for cleaning up soils which are highly polluted by heavy metals. Limiting the movement of pollutants out of the contaminated area by creating a dense and persistent plant cover appears to be the more reasonable approach. In this context, phytostabilization is a technique that uses metallicolous plants to revegetate highly polluted soils.This paper presents the results of an experiment performed in situ using metallicolous ecotypes of four plant species native to the Mediterranean French region, and grown in different combinations at a polluted site over two years. The soils were highly polluted with zinc, cadmium and lead. The aim was to find the best species mixture in terms of cover, biomass and duration. The four species used were the biennial legume Anthyllis vulneraria, two perennial grasses, Festuca arvernensis and Koeleria vallesiana, and the perennial forb Armeria arenaria. Mixtures which included A. vulneraria, and especially when in combination with F. arvernensis, showed the highest values of cover and biomass. After flowering, the biennial individuals of A. vulneraria disappeared but subsequent germination and survival of seedlings occurred abundantly under the two grasses. Mixtures with A. arenaria showed the lowest values of cover and biomass. Soil nitrogen increased in the plots with A. vulneraria as well as the concentration of essential nutrients (N P K) in the aerial parts of the two grasses. In contrast, the concentration of metals (Zn Pb Cd) decreased in the aboveground biomass of the latter in the same plots.These results show that reciprocal facilitation effects can act in heavy metal polluted environments, and that phytostabilization efforts in the Mediterranean region can be improved by using mixtures including local metallicolous legume and grass species.

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Some plant species growing on metalliferous soils are able to accumulate heavy metals in their shoots up to very high concentrations, but the selective advantage of this behaviour is still unknown. The most popular hypothesis, that metals protect plants against herbivores, has been tested several times in laboratory conditions, with contradictory results. We carried out the first large-scale test of the defence hypothesis in eight natural populations of the model Zn hyperaccumulator Thlaspi caerulescens J. and C. Presl (Brassicaceae). In two climatic regions (temperate, Belgium-Luxembourg, and Mediterranean, southern France), we worked in metalliferous and in normal, uncontaminated environments, with plants spanning a wide range of Zn concentrations. We also examined the importance of glucosinolates (main secondary metabolites of Brassicaceae) as antiherbivore defences. When exposed to natural herbivore populations, T. caerulescens suffered lower herbivory pressures in metal-enriched soils than in normal soils, both in Belgium-Luxembourg and in southern France. The trapping of gastropods shows an overall lower population density in metalliferous compared to normal environments, which suggests that herbivory pressure from gastropods is lower on metalliferous soils. In addition, foliar concentration of glucosinolates was constitutively lower in all populations from metal-enriched soils, suggesting that these have evolved towards lower investment in organic defences in response to lower herbivory pressure. The Zn concentration of plants had a protective role only for Belgian metallicolous plants when transplanted in normal soils of Luxembourg. These results do not support the hypothesis that Zn plays a key role in the protection of T. caerulescens against enemies. In contrast, glucosinolates appear to be directly involved in the defence of this hyperaccumulator against herbivores.

Several components of the diversity of plant communities, such as species richness, species composition, number of functional groups and functional composition, have been shown to directly affect the performance of exotic species. Exotics can also be affected by herbivores of the native plant community. However, these two possible mechanisms limiting invasion have never been investigated together. The aim of this study was to investigate the relationships between plant diversity, herbivory and performance of two annual exotics, Conyza bonariensis and C. canadensis, in Mediterranean annual communities. We wanted to test whether herbivory of these exotics was influenced either by species richness, functional-group richness or functional-group composition. We also studied the relationship between herbivory on the exotic species and their performance. Herbivory increased with increasing species and functional-group richness for both Conyza species. These patterns are interpreted as reflecting a greater number of available herbivore niches in a richer, more complex, plant community. The identities of functional groups also affected Conyza herbivory, which decreased in the presence of Asteraceae or Fabaceae and increased in the presence of Poaceae. Increasing herbivory had consequences for vegetative and demographic parameters of both invasive species: survival, final biomass and net fecundity decreased with increasing herbivory, leading to a loss of reproductive capacity. We conclude that communities characterised by a high number of grass species instead of Asteraceae or Fabaceae may be more resistant to invasion by the two Conyza species, in part due to predation by native herbivores.

Recent studies have shown that a high species or functional group richness may not always lead to a greater resistance of plant communities to invasion, whereas species and/or functional group composition can more reliably predict invasion resistance. The aim of this study was to understand the mechanisms through which functional group composition can influence the resistance of Mediterranean annual communities to invasion by the exotic Conyza bonariensis. To analyse the effects of functional composition on the performance of individuals introduced as seedlings we first examined the relationships between the demographic and vegetative parameters of C. bonariensis and the biomass achieved by each functional group (grasses, legumes and Asteraceae rosettes) in synthetic communities. As a further step to approach the mechanisms involved in community resistance to invasion, we included in the analyses measurements of functional variables taken within the synthetic communities. In agreement with earlier results and theory suggesting that high nutrient availability can favour invasions, an abundant legume biomass in communities increased the final biomass and net fecundity of C. bonariensis, due to positive effects on soil nitrate concentration. Survival and establishment of C. bonariensis were mainly favoured by a high biomass of Asteraceae. Additional results from measurements of herbivory suggested that C. bonariensis survival wasn't related to abiotic conditions but may be owed to a protection against herbivores in plots with abundant Asteraceae. Establishment was on the other hand likely to be hindered by the effects of abundant grass and legume foliage on light quality, and therefore easier within an Asteraceae canopy. We conclude that invasion of Mediterranean old fields by species with biologies similar to C. bonariensis could be limited by favouring communities dominated by annual grasses.

Summary The relationship between zinc (Zn) tolerance and hyperaccumulation in Thlaspi caerulescens was investigated from F1 and F2 crosses within and among metallicolous and nonmetallicolous Mediterranean populations. F1 offspring were grown on increasingly Zn‐enriched soils to test Zn enrichment effects, and many families of F2 offspring were grown on a Zn‐rich soil. Tolerance of F1 offspring depended on stress intensity. Tolerance of interecotype crosses was intermediate between that of the intraecotype crosses. No difference in Zn accumulation appeared among the F1 offspring from the three crosses involving metallicolous parents. Otherwise, none of these offspring exceeded the Zn hyperaccumulation threshold (10 000 mg kg−1), unlike the nonmetallicolous ones. The latter also showed the highest mortality. In some F2 families from interecotype crosses, hyperaccumulation values exceeded 15 000 mg kg−1 in nontolerant offspring, whereas tolerant offspring displayed lower values (c. 10 000 mg kg−1). There was no difference between tolerant and nontolerant offspring when they showed low hyperaccumulation. Therefore, the relationship between tolerance and hyperaccumulation in F1 and F2 crosses depended on the hyperaccumulation level of plants.

Although the causes and consequences of social dominance have been examined extensively, avian studies have rarely focused on between‐population differences in social dominance. On the island of Corsica, two resident blue tit Parus caeruleus populations 25 km apart differ significantly in body size measures, timing and effort of reproduction, and song structure, and some of these population differences have a genetic basis. Because earlier avian studies have shown that social dominance is influenced by body size or mass, we predicted that individuals from these two blue tit populations would also differ in their ability to dominate other individuals. Consistent with this prediction, we found that male blue tits of these two populations differ in social dominance, and that heavier or larger individuals dominate lighter or smaller ones in aviary experiments. We propose that social dominance may serve to maintain phenotypic population differentiation at a micro‐geographic scale by acting as a barrier to dispersal.