Aims Human impacts on natural ecosystems induce changes in their functioning through alterations in species richness, composition and evenness of plant communities. Most litter diversity–decomposition processes studies have only manipulated species richness, ignoring the role of evenness. Here, results from a field litterbag experiment are presented to test whether changes in evenness of species distribution in litter mixtures affected the strength of the litter-species richness–decomposition relationship.
Methods Ten herbaceous species abundant in Mediterranean grassland communities and representative of different genera and functional groups were used. Species richness was directly manipulated to produce litter mixtures of three and six plant species, as well as litter of each individual species used. Each level of species richness was replicated several times such that each repeat had a different species composition. Three- and six-species litter mixtures were also treated to vary in evenness (three levels). Decomposition rate was assessed by percentage dry weight loss over the 90 days of the experiment.
Important findings Decomposition rate was positively related to the linear increase in litter-species richness and was affected by the composition of the litter-species mixture. Decomposition rates differed significantly between evenness treatments and moreover, the strength of the positive relationship between litter-species richness and decomposition rate decreased notably in the low-evenness treatment. The effects of evenness on decomposition rate, at different richness levels, were partially explained by the differences in the initial litter mixture's carbon-to-nitrogen ratio within them. This study reveals that short-term decomposition rate is positively affected by both components of Mediterranean grassland litter-species diversity.

Aims Poplars grown in North China may experience water-deficient periods in their life cycle. The aim of the present paper was to quantify the response of three clones to different watering regimes and to determine which clone among the three is the best adapted to drought conditions.
Methods Three hybrid poplar clones (clone DN-34, R-247 and OP-367) were used in the present experiment. The seedlings of the three clones were grown under four watering regimes: control (well watered, 100% field water capacity (FC)) and three drought treatments (drought stress I, 50% FC; drought stress II, 40% FC; drought stress III, 30% FC). Changes in morphological, physical and biochemical indicators of the three hybrid poplar clones were investigated.
Important findings Drought treatment (50%, 40% and 30% FC) decreased net photosynthetic rate (Pn), transpiration rate (Tr), stomatal conductance (gs), shoot height, total biomass and chlorophyll (Chl) content in all the three clones and it increased activities of antioxidant enzymes and free proline content. The highest values of the above-mentioned morphological and physiological parameters were recorded in clone OP-367 under 30% FC, followed by clone DN-34 and R-247. Relative leaf water content (RWC) and stem diameter (sd) markedly declined in clone R-247 and DN-34 under drought stress I, II and III, whereas RWC and sd declined in clone OP-367 only under drought stress II and III. Clone OP-367 had more RWC and sd than DN-34 and R-247. Only the 30% FC induced an increase in the root-to-shoot ratio (rs) and water use efficiency (WUE) in all the three clones. OP-367 was the most efficient clone in water absorption and use, for plants of the clone had the highest values of rs and WUE. Our data demonstrate that among the three clones, OP-367 was better able to maintain photosynthesis and growth and lower the damage caused by drought.

Aims Recent studies have shown that alpine meadows on the Qinghai-Tibetan plateau act as significant CO₂ sinks. On the plateau, alpine shrub meadow is one of typical grassland ecosystems. The major alpine shrub on the plateau is Potentilla fruticosa L. (Rosaceae), which is distributed widely from 3 200 to 4 000 m. Shrub species play an important role on carbon sequestration in grassland ecosystems. In addition, alpine shrubs are sensitive to climate change such as global warming. Considering global warming, the biomass and productivity of P. fruticosa will increase on Qinghai-Tibetan Plateau. Thus, understanding the carbon dynamics in alpine shrub meadow and the role of shrubs around the upper distribution limit at present is essential to predict the change in carbon sequestration on the plateau. However, the role of shrubs on the carbon dynamics in alpine shrub meadow remains unclear. The objectives of the present study were to evaluate the magnitude of CO₂ exchange of P. fruticosa shrub patches around the upper distribution limit and to elucidate the role of P. fruticosa on ecosystem CO₂ fluxes in an alpine meadow.
Methods We used the static acrylic chamber technique to measure and estimate the net ecosystem productivity (NEP), ecosystem respiration (R e), and gross primary productivity (GPP) of P. fruticosa shrub patches at three elevations around the species' upper distribution limit. Ecosystem CO₂ fluxes and environmental factors were measured from 17 to 20 July 2008 at 3 400, 3 600, and 3 800 m a.s.l. We examined the maximum GPP at infinite light (GPP max) and maximum R e (R emax) during the experimental time at each elevation in relation to aboveground biomass and environmental factors, including air and soil temperature, and soil water content.
Important findings Patches of P. fruticosa around the species' upper distribution limit absorbed CO₂, at least during the daytime. Maximum NEP at infinite light (NEP max) and GPP max of shrub patches in the alpine meadow varied among the three elevations, with the highest values at 3 400 m and the lowest at 3 800 m. GPP max was positively correlated with the green biomass of P. fruticosa more strongly than with total green biomass, suggesting that P. fruticosa is the major contributor to CO₂ uptake in the alpine shrub meadow. Air temperature influenced the potential GPP at the shrub-patch scale. R emax was correlated with aboveground biomass and R emax normalized by aboveground biomass was influenced by soil water content. Potentilla fruticosa height (biomass) and frequency increased clearly as elevation decreased, which promotes the large-scale spatial variation of carbon uptake and the strength of the carbon sink at lower elevations.

Aims Species-rich plant communities are hypothesized to be more resistant against plant invasions because they use resources in a more efficient way. However, the relative contributions of aboveground competition and belowground interactions for invasion resistance are still poorly understood.
Methods We compared the performance of Knautia arvensis transplants growing in plots differing in plant diversity both under full competition and with shoots of neighbors tied back to determine the relative strength of aboveground competition in suppressing this test invader without the confounding effect of shading. In addition, we assessed the effects of belowground competition and soil-borne pathogens on transplant performance.
Important findings Both aboveground competition and plant species richness strongly and independently affected invader performance. Aboveground biomass, height, leaf mass per area and flowering of transplanted individuals of K. arvensis decreased with increasing species richness of the host community. Species-rich and species-poor communities both imposed equally strong aboveground competition on K. arvensis. However, belowground interactions (especially belowground root competition) had strong negative effects on transplant performance. In addition, the presence of grasses in a plant community further reduced the performance of K. arvensis. Our results suggest that belowground competition can render species-rich host communities more suppressive to newly arriving species, thus enhancing community invasion resistance.

Aims Invasion resistance in experimental plant communities is known to increase with increasing diversity and further to depend on the presence of particular functional groups. To test whether these effects also hold true for the invader establishment phase beyond the seedling stage, we studied survival and performance of Centaurea jacea L. (brown knapweed) planted into experimental grassland communities of varying plant biodiversity over three consecutive years. Moreover, we analysed the role of insect herbivory and biomass of the recipient community for mediating diversity effects.
Methods In 2005, seedlings of Centaurea were transplanted into experimental grassland communities (the Jena Experiment) covering a species richness (1–60) and functional group richness (1–4) gradient. Half of these transplants and the community surrounding them in each plot were sprayed with insecticide while the other half served as control. In 2006 and 2007 (during the second and third year after transplantation), we recorded survival, growth-related (e.g. transplant biomass, height) and reproduction-related traits (e.g. number of flower heads). Annual data on community aboveground biomass served as covariate to investigate mediating effects of aboveground competition with the recipient community.
Important findings Species richness was the most important factor responsible for Centaurea limitation. Higher levels of diversity decreased survival and all performance traits in both years. These diversity effects were partly driven by community biomass, but not fully explained by that covariate, suggesting the importance also of further processes. The influence of functional group richness was strong in the second year after transplantation and weaker in the third year. Among the particular functional groups, only the presence of legumes showed strong negative effects on Centaurea survival and weak negative effects on growth and reproduction, the latter two being mediated by biomass. Insect herbivore reduction considerably benefited Centaurea in sprayed monocultures, where it grew significantly larger than in all other diversity levels and than in the control subplots. We conclude that effects of plant community properties on invading individuals change in the course of establishment, that plant species richness effects are also important during later stages of establishment, and that biomass (especially at high diversity) and herbivory (especially at low diversity) of the recipient community are important in mediating community effects on invaders.

Aims Nitrogen (N) and phosphorus (P) are limiting nutrients to life across a variety of ecosystems. N:P stoichiometry, concerning the balance of these two elements, has recently received great attention. However, little is known about the nature of N:P stoichiometry in obligate mutualism.
Methods N:P stoichiometry of Ficus racemosa and its pollinating wasp Ceratosolen fusciceps, an example of coevolving obligate mutualism, was investigated, and the N:P stoichiometric traits of male versus female wasps were compared.
Important findings Nutrient concentrations in C. fusciceps were much higher than in its host. N enrichment in fig wasp was evidently stronger than phosphorus. N concentrations of male fig wasps were significantly higher than those of females, while P concentrations of female fig wasps were remarkably higher than those of male ones. Therefore, N:P ratios in male fig wasps were significantly greater than in female fig wasps. N:P ratio in fig-pollinating wasp displayed linear functions to fig N contents, suggesting that N limitation in fig wasps may dominate the nutritional relationship between fig pollinator and its host. Fig wasp population size had significant influences on N concentrations in host fig and female wasp per se. Driven by the nutritional stress of pollinating and parasite insects, fig fruit preferred increasing its diameter first but not nutrient richness. Values for N and P contents of fig pollinators showed seasonal differences with greater N:P ratios in dry season than in rainy season. The observations suggest that tropical climate change would result in more severe N limitation to fig-pollinating wasp and may further influence the stability of fig–fig wasp mutualism.

Aims Most plants are clonal in nature. Clonal ramets can share water, nutrients and photosynthate, especially when they experience patchy resources. Patch contrast (i.e. a difference in resources among patches) and patch direction (i.e. source–sink relations) are among the basic attributes of spatial patchiness. Here, I hypothesize that young established ramets in nutrient-rich patches support old ramets in nutrient-poor patches when ramets are subjected to different patch contrasts and patch directions.
Methods In a greenhouse experiment, old and young ramets of Glechoma longituba were grown in four combinations consisting of patch contrast and patch direction. Minus patch direction refers to a patch combination in which parent ramets grow in nutrient-rich patches while connected daughter ramets grow in nutrient-poor ones and plus patch direction is the opposite direction. I measured photosynthesis and fluorescence traits, harvested all ramets, took morphological measures, weighed their dry mass and determined their nutrient uptake and use.
Important findings For parental ramets of G. longituba, patch contrast and patch direction and their interactions had no significant effects on net photosynthetic rate, maximal fluorescence yield, photochemical quenching (quenching refers to any process which decreases the fluorescence intensity of a given substance), non-photochemical quenching, nutrient uptake, biomass and stolon weight ratio. Patch direction alone significantly affected root weight ratio. Large patch contrast enhanced N use efficiency (NUE) and P use efficiency (PUE); plus patch direction decreased NUE, but increased PUE; the patch contrast by patch direction interaction affected PUE and K use efficiency (KUE). There were significant interactions between patch direction and patch contrast on PUE and KUE. It is concluded that soil nutrient patchiness may influence nutrient use strategies, but not nutrient uptake, photosynthesis and growth of parent ramets of G. longituba connected to daughter ramets, and that patch contrast and patch direction jointly affect PUE and KUE.

Aims Coastal areas, and in particular coastal dunes, are ecosystems strongly affected by the invasion of alien plants. However, few attempts have ever been made to quantify alien species incidence in different communities along the coastal zonation. This work aims to analyze the distribution of alien plants along the coastal zonation of sandy shores on the Tyrrhenian coast, addressing specifically differences among plant communities in abundance of alien plants.
Methods The study was performed on recent dunes (Holocene) of the central western coast of Italy. We selected dune landscapes where invasion processes were particularly evident. Vegetation plots were randomly sampled and through cluster analysis, we identified six plant communities corresponding to the typical zonation described for the Tyrrhenian sandy coast of Central Italy. We evaluated and compared frequency and abundance of invasion in these different communities. Further, we investigated how propagule pressure (measured using as proxy human structures) contributed to the observed invasion patterns.
Important findings We found a relatively low total number of aliens but also a differential distribution pattern and strong abundance of some of the aliens in specific sectors of the vegetation zonation. The perennial community of transition dunes appears most affected by invasion processes, related almost exclusively to the frequent and widespread Carpobrotus aff. acinaciformis. This alien species reaches high cover values, apparently lowering cover of native species of transition dune plant communities. Higher levels of invasion in the transition dune can be partially explained because of greater propagule pressure in this section of the dune profile. Our findings thus have important conservation and management implications since transition dune communities with Crucianella maritima are rare and protected (sensu Habitat 92/43/EEC Directive) along the entire Italian coast.