Hydraulic lift and tolerance to salinity of semiarid species: consequences for species interactions

The different abilities of plant species to use ephemeral or permanent water sources strongly affect physiological performance and species coexistence in water-limited ecosystems. In addition to withstanding drought, plants in coastal habitats often have to withstand highly saline soils, an additional ecological stress. Here we tested whether observed competitive abilities and C–water relations of two interacting shrub species from an arid coastal system were more related to differences in root architecture or salinity tolerance. We explored water sources of interacting Juniperus phoenicea Guss. and Pistacia lentiscus L. plants by conducting physiology measurements, including water relations, CO₂ exchange, photochemical efficiency, sap osmolality, and water and C isotopes. We also conducted parallel soil analyses that included electrical conductivity, humidity, and water isotopes. During drought, Pistacia shrubs relied primarily on permanent salty groundwater, while isolated Juniperus plants took up the scarce and relatively fresh water stored in upper soil layers. As drought progressed further, the physiological activity of Juniperus plants nearly stopped while Pistacia plants were only slightly affected. Juniperus plants growing with Pistacia had stem-water isotopes that matched Pistacia, unlike values for isolated Juniperus plants. This result suggests that Pistacia shrubs supplied water to nearby Juniperus plants through hydraulic lift. This lifted water, however, did not appear to benefit Juniperus plants, as their physiological performance with co-occurring Pistacia plants was poor, including lower water potentials and rates of photosynthesis than isolated plants. Juniperus was more salt sensitive than Pistacia, which withstood salinity levels similar to that of groundwater. Overall, the different abilities of the two species to use salty water appear to drive the outcome of their interaction, resulting in asymmetric competition where Juniperus is negatively affected by Pistacia. Salt also seems to mediate the interaction between the two species, negating the potential positive effects of an additional water source via hydraulic lift.     

Ecophysiological significance of chlorophyll loss and reduced photochemical efficiency under extreme aridity in Stipa tenacissima L.

Stipa tenacissima L., a perennial tussock grass widely found in semi-arid environments of the Iberian Peninsula and North Africa, is subjected to multiple stresses during the extreme summer conditions of south-east Spain. We characterised the photoprotective mechanisms of S. tenacissima during the transition from spring to summer and autumn. S. tenacissima experienced a marked water deficit (_{{ pd}} < -8.4 MPa) and the complete suppression of CO₂ assimilation in August, associated with a 72% reduction of maximal photochemical efficiency of PSII (F_{{ v}}/F_{{ m}}). These reduced F _{{ v}}/F_{{ m}} values were related to the pre-dawn maintenance of high levels of epoxidized forms of xanthophyll-cycle pigments (DPS_{{ pd}}, ca. 42% higher than spring values), and with a 60% reduction in the concentration of total chlorophyll (Chl a+b). These changes were associated with a low capacity of dissipation of the excitation energy non-radiatively (measured as NPQ). Leaves showed a complete recovery of F _{{ v}}/F_{{ m}} and xanthophyll and chlorophyll concentrations after the autumn rainfall, which reached levels similar to that of spring. This poikilohydric-type response of S. tenacissima to stress allows for a greater tolerance of water shortage, high temperature and high light intensity, which are typical in these semi-arid environments and accounts for its distinctive opportunistic growth.     

Phenological and reproductive responses of a semiarid shrub to pulsed watering

Climate models forecast the decrease in annual precipitation and more extreme events (i.e. fewer rainy days and longer drought periods between rainfall events) in Mediterranean ecosystems which may have important consequences for plant community dynamics. Water availability is the most limiting factor for plant performance in arid environments, where it is often highly pulsed, and discrete rainfall events interspersed with drought periods are the important components of the annual water supply. Therefore, relatively small changes in rainfall frequency can have strong effects on the plant communities. We tested the effects of fewer rain events on inflorescence and fruit phenology, and reproductive success of a Mediterranean shrub species from semiarid SE Spain in a glasshouse trial. We found that by changing patterns of water supply, we strongly altered soil drying cycles with consequences for Phagnalon saxatile life history. Inflorescence and fruit phenology were not affected, and no changes in the number of inflorescences, fruits or fruit set were recorded. However, individuals growing under lower rainfall frequency produced bigger shoots, and heavier fruits and seeds. Bigger fruits produced a higher number of seeds that were heavier and more viable and, therefore, produced more seedlings than plants growing in the control treatment. Higher reproductive success achieved by Phagnalon individuals under low watering frequency may have important consequences for community dynamics if larger but less frequent rain events lead to an increase in soil water availability: natural selection favours individuals making the greatest contribution to the next generation. Such advantages could change community composition, the spatial distribution of species and their interactions.     

Do biotic interactions shape both sides of the humped‐back model of species richness in plant communities?

A humped-back relationship between species richness and community biomass has frequently been observed in plant communities, at both local and regional scales, although often improperly called a productivity-diversity relationship. Explanations for this relationship have emphasized the role of competitive exclusion, probably because at the time when the relationship was first examined, competition was considered to be the significant biotic filter structuring plant communities. However, over the last 15 years there has been a renewed interest in facilitation and this research has shown a clear link between the role of facilitation in structuring communities and both community biomass and the severity of the environment. Although facilitation may enlarge the realized niche of species and increase community richness in stressful environments, there has only been one previous attempt to revisit the humped-back model of species richness and to include facilitative processes. However, to date, no model has explored whether biotic interactions can potentially shape both sides of the humped-back model for species richness commonly detected in plant communities. Here, we propose a revision of Grime's original model that incorporates a new understanding of the role of facilitative interactions in plant communities. In this revised model, facilitation promotes diversity at medium to high environmental severity levels, by expanding the realized niche of stress-intolerant competitive species into harsh physical conditions. However, when environmental conditions become extremely severe the positive effects of the benefactors wane (as supported by recent research on facilitative interactions in extremely severe environments) and diversity is reduced. Conversely, with decreasing stress along the biomass gradient, facilitation decreases because stress-intolerant species become able to exist away from the canopy of the stress-tolerant species (as proposed by facilitation theory). At the same time competition increases for stress-tolerant species, reducing diversity in the most benign conditions (as proposed by models of competition theory). In this way our inclusion of facilitation into the classic model of plant species diversity and community biomass generates a more powerful and richer predictive framework for understanding the role of plant interactions in changing diversity. We then use our revised model to explain both the observed discrepancies between natural patterns of species richness and community biomass and the results of experimental studies of the impact of biodiversity on the productivity of herbaceous communities. It is clear that explicit consideration of concurrent changes in stress-tolerant and competitive species enhances our capacity to explain and interpret patterns in plant community diversity with respect to environmental severity.     

Is the cask of facilitation ready for bottling? A symposium on the connectivity and future directions of positive plant interactions

The 2009 British Ecological Society''s Annual Symposium entitled ‘Facilitation in Plant Communities’ was held at the University of Aberdeen, Scotland, from 20 to 22 April 2009. This was the first ever international meeting dedicated to the rapidly expanding field of facilitation. The aim of the symposium was to assess the current ‘state-of-play’ by contrasting findings from different systems and by looking outwards in an attempt to integrate this field with other related fields. It was also aimed at understanding how knowledge of facilitation can help understand community dynamics and be applied to ecosystem restoration. The symposium identified several key areas where future work is likely to be most profitable.