Ivan Nijs

Can complementarity in water use help to explain diversity–productivity relationships in experimental grassland plots?

Positive diversity–productivity relationships have repeatedly been found in experimental grassland plots, but mechanistic explanations are still under debate. We tested whether complementarity for the exploitation of the soil water resource helps to explain these relationships. In the dry summer of 2003, evapotranspiration (ET) was assessed at the Swedish BIODEPTH site using two different approaches: snapshot measurements of canopy surface temperature and simulation of time-accumulated ET by means of a soil water balance model. More diverse plots were characterized by lower surface temperatures and higher accumulated ET. Transgressive overyielding tests revealed that ET in polycultures was higher than in the best-performing monocultures, but this pattern was reversed at high degrees of water stress. Our results indicate that a more complete exploitation of soil water by more diverse grassland systems is on the one hand likely to be a driver for their increased biomass production, but on the other hand causes the more diverse communities to be affected earlier by drought. Nevertheless, the results also suggest that productivity may (at least partially) be maintained under dry conditions due to the higher likelihood of including drought-tolerant species in the more diverse communities.

Survival, Gap Formation, and Recovery Dynamics in Grassland Ecosystems Exposed to Heat Extremes: The Role of Species Richness

A field experiment was performed in which the richness of perennial grasses (S) was varied in model ecosystems exposed to a simulated heat wave (free air temperature increase and drought). The proportion of individuals that survived the heat wave decreased with S, which could be ascribed to higher water consumption in the species-rich systems. Higher transpiration at high diversity was also observed in other studies using functional groups and could have originated from increased leaf area, less intense stomatal closure, or a combination of both. The increased tiller number per plant that we observed, while leaf area per tiller remained constant, suggests that an enhanced leaf area index was most likely responsible. However, competitive interactions also seemed to play a role in the influence of S on survival. Regrowth of the surviving individuals, expressed as leaf area per living plant after a recovery period following the heat wave, increased with S, most likely due to the dominance of productive species, which was facilitated by the additional space yielded by more intense gap formation at higher S (due to higher plant mortality). Species richness affected both the size and density of the gaps. Mean size increased exponentially with S, while density increased at low S but decreased at higher S when connectance of the gaps occurred. Size distribution of the gaps was not affected.

Factors determining plant–neighbour interactions on different spatial scales in young species-rich grassland communities

In naturally colonised species-rich grassland communities, we examined the properties of a plant’s aboveground neighbourhood that affect its performance (aboveground biomass). To this end a range of neighbourhood parameters were measured: number, biomass and species richness of the neighbours, number and biomass of the conspecific neighbours, and light availability at the base of the target plant. We also determined at which neighbourhood size the strongest target plant–neighbour interactions occurred, and whether conspecific neighbours affected competitively stronger or weaker target species differently. Target plant performance varied with target identity, and was significantly affected by light availability and the number of neighbouring plants (neighbourhood density). Depending on the target species, there was also an effect of total neighbour biomass on plant performance. The target plants were most strongly affected by their neighbours within a 3-cm distance, which could account for 78% of the variance in target biomass. Number or biomass of the conspecific neighbours did not contribute to the explanation of target performance in any of the target species. Whereas in an 8-cm neighbourhood the amount of light penetration was the strongest predictor of target performance, the number of neighbours was more important in a 3-cm neighbourhood. These experimental results might be useful to extend existing neighbourhood competition models for one or two species to multi-species competition models.

MORPHOLOGICAL AND PHYSIOLOGICAL INDICATORS OF TOLERANCE TO ATMOSPHERIC STRESS IN TWO SENSITIVE AND TWO TOLERANT TEA CLONES IN SOUTH AFRICA

Tea ( Camellia sinensis ) clones (PC113 and SFS204) sensitive to very dry air and clones (PC114 and SFS150) that are tolerant, were studied at two tea estates (Tshivhase and Grenshoek) in the Northern Province of the Republic of South Africa. Among the morphological leaf traits studied, stomatal density, pore diameter and pore depth were not linked consistently to stress tolerance. Cuticle thickness was not a good indicator of stress tolerance because genetic differences between clones were confounded by the clonal response of wax production to stress. In contrast, measured leaf conductance to water vapour transport was larger and leaf water potential was lower in sensitive clones, but only with more severe atmospheric stress (Grenshoek). Also the ratio of the calculated maximum stomatal conductance in old and young leaves was higher in sensitive clones, suggesting that the loss of a larger fraction of the total stem flow by old leaves enhanced the stress experienced by the young leaves. However, this indicator was valid only under the more stressful microclimate of Grenshoek. The results indicate that even promising criteria for stress tolerance should be tested by exposure to stress during selection.

Dry matter yield and herbage quality of field margin vegetation as a function of vegetation development and management regime

Dry matter (DM) yield and herbage quality of unfertilized mown field margin strips were studied during early succession in a field experiment over a period of three years. The experiment aimed to maximize botanical diversity and was conducted at two different locations with contrasting soil type and comprised four vegetation types (spontaneously regenerated versus sown vegetation) and three herbage removal strategies (herbage left versus herbage removed). The experimental factors investigated were location, vegetation and herbage removal. Margin strips were mown twice a year with a late first cut around 15 June and a regrowth cut around 15 September to meet nature conservation objectives. Average DM yield over the first three years was not significantly affected by herbage removal but increased significantly over time, irrespective of vegetation or herbage removal. Initially, sown margin strips significantly outyielded unsown margin strips, but differences in DM yield converged over time. The mid-June cut yielded significantly more than the regrowth cut but its herbage quality was significantly lower. Herbage from the unsown margin strip had a significantly better forage quality than herbage from sown margin strips. Forage quality decreased over time, irrespective of location or vegetation. Changes over time in DM yield and quality were attributed to changes in species composition. The herbage quality of field margins was lower than the herbage quality of intensively managed grassland, limiting its use in rations for highly productive livestock.

Robustness of metacommunities with omnivory to habitat destruction: disentangling patch fragmentation from patch loss

Habitat destruction, characterized by patch loss and fragmentation, is a major driving force of species extinction, and understanding its mechanisms has become a central issue in biodiversity conservation. Numerous studies have explored the effect of patch loss on food web dynamics, but ignored the critical role of patch fragmentation. Here we develop an extended patch-dynamic model for a tri-trophic omnivory system with trophic-dependent dispersal in fragmented landscapes. We found that species display different vulnerabilities to both patch loss and fragmentation, depending on their dispersal range and trophic position. The resulting trophic structure varies depending on the degree of habitat loss and fragmentation, due to a tradeoff between bottom-up control on omnivores (dominated by patch loss) and dispersal limitation on intermediate consumers (dominated by patch fragmentation). Overall, we find that omnivory increases system robustness to habitat destruction relative to a simple food chain.

Combined elevated CO2 and climate warming induces lagged effects of drought in Lolium perenne and Plantago lanceolata

Future climate scenarios predict increases in elevated atmospheric CO2, air temperature and drought, but the impacts of multiple climate change factors on ecosystem functioning remain unclear. In this study, we compared drought responses of plants under future versus current climate conditions. In addition to focusing on stress during the drought itself, we also examined post-drought lagged effects, and whether warming and elevated CO2 alter these. We grew monocultures and mixtures of two grassland species (Lolium perenne L. and Plantago lanceolata L.) in four simulated climate scenarios: (1) current climate, (2) current climate with drought, (3) warmer temperature with drought and (4) combined warming, elevated CO2 and drought. L. perenne and P. lanceolata were influenced by the climate scenario but not differently enough to modify the competitive balance. Warming aggravated drought impacts on L. perenne and elevated CO2 only partly compensated for these effects. In a warmer climate, with or without elevated CO2, drought continued to enhance senescence and mortality in L. perenne long after the water shortage, while such lag effects were not observed in current climate. In P. lanceolata, a similar stimulation of senescence and mortality was induced, but only under combined warming and elevated CO2. These lag effects induced by the future climate may reduce resilience.