Questions: Is there any evidence of coordination among leaf, stem and root traits, and thereby of the existence of a plant economics spectrum at the species and community level in Mediterranean forests? Are these traits related to plant size and seed mass?Location: Mediterranean forests and shrublands, Sierra Morena mountains, C ordoba, southern Spain.
Methods:We selected nine woody plant communities along a natural local gradient of soil water and nutrient availability. We measured key leaf, stem, root and whole-plant traits for 38 dominant woody plant species. The variation across species of 15 functional traits (of the leaf, stem and root) was analysed and coordination among them was tested. We explored the relationships between these traits (hereafter 'resource-use traits' due to their close association with the acquisition-conservation trade-off) and plant height and seed mass. Finally, we compared results at species level with those calculated at community level, considering community-weighted means (CWMs).
Results:We found a significant coordination between traits belonging to different plant organs, and propose the existence of a plant economics spectrum in Mediterranean forests along the environmental gradient. However, weaker relationships were found within groups of species under similar environmental conditions. We did not find the expected orthogonal relationships between plant height, seed mass and resource-use traits. Relationships among functional traits were stronger at the community level than at the species level.Conclusions: This study reveals a high degree of functional coordination between traits belonging to different plant organs at both species and community level, and suggests the existence of a plant economics spectrum across 38 Mediterranean woody plant species. However, this general trend of functional coordination between organs became weaker or disappeared when considering restricted groups of species belonging to environmentally similar sites (e.g. dry vs wet sites), suggesting that the diversification of strategies within communities is not related to the economics spectrum at a lower spatial scale. Interestingly, the high degree of coordination between resource-use traits and seed mass at the community level seems to support the tolerance-fecundity model, which predicts an inverse relationship between fecundity and stress tolerance.
Leaf mass per area (LMA) is a morphological trait widely used as a good indicator of plant functioning (i.e. photosynthetic and respiratory rates, chemical composition, resistance to herbivory, etc.). The LMA can be broken down into the leaf density (LD) and leaf volume to area ratio (LVA or thickness), which in turn are determined by anatomical tissues and chemical composition. The aim of this study is to understand the anatomical and chemical characteristics related to LMA variation in species growing in the field along a water availability gradient. We determined LMA and its components (LD, LVA and anatomical tissues) for 34 Mediterranean (20 evergreen and 14 deciduous) woody species. Variation in LMA was due to variation in both LD and LVA. For both deciduous and evergreen species LVA variation was strongly and positively related with mesophyll volume per area (VA or thickness), but for evergreen species positive relationships of LVA with the VA of epidermis, vascular plus sclerenchyma tissues and air spaces were found as well. The leaf carbon concentration was positively related with mesophyll VA in deciduous species, and with VA of vascular plus sclerenchymatic tissues in evergreens. Species occurring at the sites with lower water availability were generally characterised by a high LMA and LD.
Understanding which factors and rules govern the process of assembly in communities constitutes one of the main challenges of plant community ecology. The presence of certain functional strategies along broad environmental gradients can help to understand the patterns observed in community assembly and the filtering mechanisms that take place. We used a trait‐based approach, quantifying variations in aboveground (leaf and stem) and belowground (root) functional traits along environmental gradients in Mediterranean forest communities (south Spain). We proposed a new practical method to quantify the relative importance of species turnover (distinguishing between species occurrence and abundance) versus intraspecific variation, which allowed us to better understand the assemblage rules of these plant communities along environmental gradients. Our results showed that the functional structure of the studied plant communities was highly determined by soil environment. Results from our modelling approach based on maximum likelihood estimators showed a predominant influence of soil water storage on most of the community functional traits. We found that changes in community functional structure along environmental gradients were mainly promoted by species turnover rather than by intraspecific variability. Specifically, our new method of variance decomposition demonstrated that between‐site trait variation was the result of changes in species occurrence rather than in the abundance of certain dominant species. In conclusion, this study showed that water availability promoted the predominance of specific trait values (both in above and belowground fractions) associated to a resource acquisition or conservation strategy. In addition, we provided evidence that changes on community functional structure along the environmental gradient were mainly promoted by a process of species replacement, which represent a crucial step towards a more general understanding of the relative importance of intraspecific versus interspecific trait variation in these woody Mediterranean communities.
Climate models predict a further drying of the Mediterranean summer. One way for plant species to persist during such climate changes is through acclimation. Here, we determine the extent to which trait plasticity in response to drought differs between species and between sites, and address the question whether there is a trade-off between drought survival and phenotypic plasticity. Throughout the summer we measured physiological traits (photosynthesis - A , stomatal conductance - g , transpiration - E, leaf water potential - ψl) and structural traits (specific leaf area - SLA, leaf density - LD, leaf dry matter content - LDMC, leaf relative water content - LRWC) of leaves of eight woody species in two sites with slightly different microclimate (north- versus south-facing slopes) in southern Spain. Plant recovery and survival was estimated after the summer drought period. We found high trait variability between species. In most variables, phenotypic plasticity was lower in the drier site. Phenotypic plasticity of SLA and LDMC correlated negatively with drought survival, which suggests a trade-off between them. On the other hand, high phenotypic plasticity of SLA and LDMC was positively related to traits associated with rapid recovery and growth after the drought period. Although phenotypic plasticity is generally seen as favourable during stress conditions, here it seemed beneficial for favourable conditions. We propose that in environments with fluctuating drought periods there can be a trade-off between drought survival and growth during favourable conditions. When climate become drier, species with high drought survival but low phenotypic plasticity might be selected for.
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