Search

The effects of altered nutrient supplies and herbivore density on species diversity vary with spatial scale, because coexistence mechanisms are scale dependent. This scale dependence may alter the shape of the species–area relationship (SAR), which can be described by changes in species richness (S) as a power function of the sample area (A): S = cAz, where c and z are constants. We analysed the effects of experimental manipulations of nutrient supply and herbivore density on species richness across a range of scales (0.01–75 m²) at 30 grasslands in 10 countries. We found that nutrient addition reduced the number of species that could co-occur locally, indicated by the SAR intercepts (log c), but did not affect the SAR slopes (z). As a result, proportional species loss due to nutrient enrichment was largely unchanged across sampling scales, whereas total species loss increased over threefold across our range of sampling scales.

The 'two-water-worlds' hypothesis is based on stable isotope differences in stream, soil and xylem waters in dual isotope space. It postulates no connectivity between bound and mobile soil waters, and preferential plant water uptake of bound soil water sources. We tested the pool-weighted impact of isotopically distinct water pools for hydrological cycling, the influence of species-specific water use and the degree of ecohydrological separation. We combined stable isotope analysis (δ18 O and δ2 H) of ecosystem water pools of precipitation, groundwater, soil and xylem water of two distinct species (Quercus suber, Cistus ladanifer) with observations of soil water contents and sap flow. Shallow soil water was evaporatively enriched during dry-down periods, but enrichment faded strongly with depth and upon precipitation events. Despite clearly distinct water sources and water-use strategies, both species displayed a highly opportunistic pattern of root water uptake. Here we offer an alternative explanation, showing that the isotopic differences between soil and plant water vs groundwater can be fully explained by spatio-temporal dynamics. Pool weighting the contribution of evaporatively enriched soil water reveals only minor annual impacts of these sources to ecosystem water cycling (c. 11% of bulk soil water and c. 14% of transpired water).

Summary 1. Tree recruitment in Mediterranean ecosystems is strongly limited at the seedling stage by drought. Increasing evidence shows the critical positive role of the canopy nurse effect on seedling survival which results from direct and indirect, positive and negative interactions between species. 2. Most studies, however, have only focused on the effects of tree canopy on water and light, ignoring other critical factors affecting seedling regeneration, such as canopy effects on high temperatures and the competing herb biomass. 3. Here, we evaluate how tree canopy cover and removal of herbs affect the survival and growth of seedlings of two dominant Mediterranean Quercus species during a 3-year study. We use an integrated model that combines several data sets to quantify and predict regeneration dynamics along environmental gradients of soil moisture, temperature and light. 4. Low soil moisture, increased soil temperature and herb biomass negatively affected seedling survival of both Quercus species. Seedling growth was positively associated with increasing soil moisture and light. 5. Although tree canopy cover directly facilitated seedling survival in both Quercus species, it also negatively affected herb biomass and thus indirectly facilitated the survival of Quercus suber, but not of Quercus ilex seedlings at low levels of soil moisture. 6. Overall, tree canopies increased seedling survival but not growth during the establishment phase, mainly by ameliorating the effects of low soil moisture and high temperatures. Tree canopy indirectly facilitated survival of Q. suber seedlings by negatively affecting the competing herb layer. 7. Synthesis and applications. To improve tree recruitment and conserve Mediterranean Quercus woodlands, the removal of herbs should be integrated into management plans for dry habitats. Interactions between abiotic and biotic factors may also effect the regeneration of these tree species. In particular, a healthy tree canopy will become important for providing conditions to facilitate seedling establishment if these habitats become drier and warmer, as predicted by some climate change scenarios.

Water is one of the major environmental factors limiting plant growth and survival in the Mediterranean region. Quercus suber L. woodlands occupy vast areas in the Iberian Peninsula, frequently under shallow water table conditions. The relative magnitude of soil and groundwater uptake to supply transpiration is not easy to evaluate under these circumstances. We recently developed a conceptual framework for the functioning of the root system in Q. suber that simulates well tree transpiration, based on two types of root behaviour: shallow connected and deep connected. Although this significantly improved knowledge on the functional traits of Mediterranean Q. suber, the approach has the limitation of requiring root sap flow data, which are seldom available. In this work, we present alternative methodologies to assess if trees are connected to groundwater and to estimate the soil and groundwater contributions to tree transpiration. We provide evidence on the tree unrestricted access to groundwater solely based on meteorological, stem sap flow and leaf water potential data. Using a soil mass balance approach, we estimated the yearly soil and groundwater contributions to tree transpiration: 69.7% and 30.3%, respectively. Groundwater uptake became dominant in the dry summer: 73.2% of tree transpiration. Results reproduce extremely well those derived from root modelling. Because of its simplicity both in formulation and data requirements, our approach is potentially liable to be adapted to other groundwater-dependent Mediterranean oak sites, where interactions between land use and water resources may be relevant. Copyright © 2013 John Wiley & Sons, Ltd.

We examined functional coordination among stem and root vulnerability to xylem cavitation, plant water transport characteristics and leaf traits in 14 co-occurring temperate tree species. Relationships were evaluated using both tra- ditional cross-species correlations and phylogenetically independent contrast (PIC) correlations. For stems, the xylem tension at which 50% of hydraulic conductivity was lost ( Ψ 50 ) was positively associated ( P < 0.001) with specific conductivity ( K S ) and with mean hydraulically weighted xylem conduit diameter ( D h-w ), but was only marginally ( P = = = 0.06) associated with leaf specific conductivity ( K L ). The PIC correlation for each of these relationships, how- ever, was not statistically significant. There was also no relationship between root Ψ Ψ Ψ 50 and root K S in either cross- species or PIC analysis. Photosynthetic rate ( A ) and stomatal conductance ( g s ) were strongly and positively correlated with root Ψ Ψ Ψ 50 in the cross-species analysis ( P < < < 0.001), a relationship that was robust to phylogenetic correction ( P < 0.01). A and g s were also positively corre- lated with stem Ψ Ψ Ψ 50 in the cross-species analysis ( P = 0.02 and 0.10, respectively). However, only A was associated with stem Ψ Ψ Ψ 50 in the PIC analysis ( P = = = 0.04). Although the relationship between vulnerability to cavitation and xylem conductivity traits within specific organs (i.e. stems and roots) was weak, the strong correlation between g s and root Ψ Ψ Ψ 50 across species suggests that there is a trade-off between vulnerability to cavitation and water transport capacity at the whole-plant level. Our results were therefore consistent with the expectation of coordination between vulnerability to xylem cavitation and the regulation of stomatal conduc- tance, and highlight the potential physiological and evolu- tionary significance of root hydraulic properties in controlling interspecific variation in leaf function.

We studied the temporal variability and resistance to perturbation of the biomass production of grassland communities from an experimental diversity gradient (the Portuguese BIODEPTH project site). With increasing species richness relative temporal variability (CV) of plant populations increased but that of communities decreased, supporting the insurance hypothesis and related theory. Species-rich communities were more productive than species-poor communities in all three years although a natural climatic perturbation in the third year (frequent frost and low precipitation) caused an overall decrease in biomass production. Resistance to this perturbation was constant across the experimental species richness gradient in relative terms, supporting a similar response from the Swiss BIODEPTH experiment. The positive biomass response was generated by different combinations of the complementarity and selection effects in different years. Complementarity effects were positive across mixtures on average in all three years and positively related to diversity in one season. The complementarity effect declined following perturbation in line with total biomass but, counter to predictions, in relative terms overyielding was maintained in all years. Selection effects were positively related to diversity in one year and negative overall in the other two years. The response to perturbation varied among species and for the same species growing in monoculture and mixture, but following the frost communities were more strongly dominated by species with lower monoculture biomass and the selection effect was more negative. In total, our results support previous findings of a positive relationship between diversity and productivity and between diversity and the temporal stability of production, but of no effect of diversity on the resistance to perturbation. We demonstrate for the first time that the relative strength of overyielding remained constant during an exceptional natural environmental perturbation.

We present a multisite analysis of the relationship between plant diversity and ecosystem functioning within the European BIODEPTH network of plant-diversity manipulation experiments. We report results of the analysis of 11 variables addressing several aspects of key ecosystem processes like biomass production, resource use (space, light, and nitrogen), and decomposition, measured across three years in plots of varying plant species richness at eight different European grassland field sites. Differences among sites explained substantial and significant amounts of the variation of most of the ecosystem processes examined. However, against this background of geographic variation, all the aspects of plant diversity and composition we examined (i.e., both numbers and types of species and functional groups) produced significant, mostly positive impacts on ecosystem processes. Analyses using the additive partitioning method revealed that complementarity effects (greater net yields than predicted from monocultures due to resource partitioning, positive interactions, etc.) were stronger and more consistent than selection effects (the covariance between monoculture yield and change in yield in mixtures) caused by dominance of species with particular traits. In general, communities with a higher diversity of species and functional groups were more productive and utilized resources more completely by intercepting more light, taking up more nitrogen, and occupying more of the available space. Diversity had significant effects through both increased vegetation cover and greater nitrogen retention by plants when this resource was more abundant through N2 fixation by legumes. However, additional positive diversity effects remained even after controlling for differences in vegetation cover and for the presence of legumes in communities. Diversity effects were stronger on above- than belowground processes. In particular, clear diversity effects on decomposition were only observed at one of the eight sites. The ecosystem effects of plant diversity also varied between sites and years. In general, diversity effects were lowest in the first year and stronger later in the experiment, indicating that they were not transitional due to community establishment. These analyses of our complete ecosystem process data set largely reinforce our previous results, and those from comparable biodiversity experiments, and extend the generality of diversity–ecosystem functioning relationships to multiple sites, years, and processes.

To investigate how plant diversity loss affects nitrogen accumulation in above-ground plant biomass and how consistent patterns are across sites of different climatic and soil conditions, we varied the number of plant species and functional groups (grasses, herbs and legumes) in experimental grassland communities across seven European experimental sites (Switzerland, Germany, Ireland, United Kingdom (Silwood Park), Portugal, Sweden and Greece). Nitrogen pools were significantly affected by both plant diversity and community composition. Two years after sowing, nitrogen pools in Germany and Switzerland strongly increased in the presence of legumes. Legume effects on nitrogen pools were less pronounced at the Swedish, Irish and Portuguese site. In Greece and UK there were no legume effects. Nitrogen concentration in total above-ground biomass was quite invariable at 1.66 ± 0.03% across all sites and diversity treatments. Thus, the presence of legumes had a positive effect on nitrogen pools by significantly increasing above-ground biomass, i.e. by increases in vegetation quantity rather than quality. At the German site with the strongest legume effect on nitrogen pools and biomass, nitrogen that was fixed symbiotically by legumes was transferred to the other plant functional groups (grasses and herbs) but varied depending on the particular legume species fixing N and the non-legume species taking it up. Nitrogen-fixation by legumes therefore appeared to be one of the major functional traits of species that influenced nitrogen accumulation and biomass production, although effects varied among sites and legume species. This study demonstrates that the consequences of species loss on the nitrogen budget of plant communities may be more severe if legume species are lost. However, our data indicate that legume species differ in their N 2 fixation. Therefore, loss of an efficient N 2 -fixer (Trifolium in our study) may have a greater influence on the ecosystem function than loss of a less efficient species (Lotus in our study). Furthermore, there is indication that P availability in the soil facilitates the legume effect on biomass production and biomass nitrogen accumulation.

Geographic variation can lead to the evolution of different local varieties, even in widespread forage plants. We investigated the performance of common forage plants in relation to their genetic diversity and local adaptation at a continental scale using reciprocal transplants at eight field sites across Europe over a 2-year period. The overall performance of the three test species, Trifolium pratense, Dactylis glomerata, Plantago lanceolata, was generally highest for plants replanted at their home site and declined with increasing transplanting distance. The three species differed in the fitness components responsible for the increased overall performance and selection advantage at home sites. In addition to the effects of local adaptation, the majority of measured traits in all three species also showed ecotypic variation. However, no single ecotype of any species was able to outperform the locally adapted strains and do best at all sites, highlighting the importance of maintaining these plant genetic resources.

We investigated the role of water use in a Mediterranean grassland, in which diversity was experimentally manipulated, and a positive relationship was observed between plant species richness and productivity. Soil moisture patterns and stable carbon isotope ratios (d 13 C) in leaves indicated greater water use by plants growing in species-rich mixtures compared to monocultures. These results suggest that complementarity or facilitation may be the mechanism responsible for the positive relationship between plant diversity and ecosystem processes.