Climate warming and plant species richness loss have been the subject of numerous experiments, but studies on their combined impact are lacking. Here we studied how both warming and species richness loss affect water use in grasslands, while identifying interactions between these global changes. Experimental ecosystems containing one, three or nine grassland species from three functional groups were grown in 12 sunlit, climate-controlled chambers (2.25 m 2 ground area) in Wilrijk, Belgium. Half of these chambers were exposed to ambient air temperatures (unheated), while the other half were warmed by 3°C (heated). Equal amounts of water were added to heated and unheated communities, so that warming would imply drier soils if evapotranspiration (ET) was higher. After an initial ET increase in response to warming, stomatal regulation and lower above-ground productivity resulted in ET values comparable with those recorded in the unheated communities. As a result of the decreased biomass production, water use efficiency (WUE) was reduced by warming. Higher complementarity and the improved competitive success of water-efficient species in mixtures led to an increased WUE in multi-species communities as compared to monocultures, regardless of the induced warming. However, since the WUE of individual species was affected in different ways by higher temperatures, compositional changes in mixtures seem likely under climatic change due to shifts in competitiveness. In conclusion, while increased complementarity and selection of water-efficient species ensured more efficient water use in mixtures than monocultures, global warming will likely decrease this WUE, and this may be most pronounced in species-rich communities.

Here we report on the single and combined impacts of climate warming and species richness on the biomass production in experimental grassland communities. Projections of a future warmer climate have stimulated studies on the response of terrestrial ecosystems to this global change. Experiments have likewise addressed the importance of species numbers for ecosystem functioning. There is, however, little knowledge on the interplay between warming and species richness. During three years, we grew experimental plant communities containing one, three or nine grassland species in 12 sunlit, climate-controlled chambers in Wilrijk, Belgium. Half of these chambers were exposed to ambient air temperatures (unheated), while the other half were warmed by 3 • C (heated). Equal amounts of water were added to heated and unheated communities, so that warming would imply drier soils if evapotranspiration was higher. Biomass production was decreased due to warming, both aboveground (-29%) and belowground (-25%), as negative impacts of increased heat and drought stress in summer prevailed. Complementarity effects, likely mostly through both increased aboveground spatial complementarity and fa-cilitative effects of legumes, led to higher shoot and root biomass in multi-species communities, regardless of the induced warming. Surprisingly, warming suppressed productivity the most in 9-species communities, which may be attributed to negative impacts of intense interspecific competi-Correspondence to: H. J. De Boeck (hans.deboeck@ua.ac.be) tion for resources under conditions of high abiotic stress. Our results suggest that warming and the associated soil drying could reduce primary production in many temperate grasslands , and that this will not necessarily be mitigated by efforts to maintain or increase species richness.

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Global warming accelerates land surface drying, increasing the incidence of extreme climatic events such as severe droughts with detrimental effects on ecosystem functions and structure. We investigated the effects of an imposed severe drought (24 days) on fully established synthesized grassland communities with three species richness (S) levels (one, three or nine species), grown for 3 years at either ambient air temperatures (unheated) or ambient +3°C (heated). Since water supply during these 3 years was equal in all treatments, heated communities experienced more frequent, short mild droughts, but it was unknown whether this conferred greater or smaller resistance for facing prolonged droughts. During the 24-day drought period, soil matric potential decreased in a similar way over time in both temperature treatments and was lower at increasing S-levels. Although green vegetation cover was significantly reduced by the drought in both temperature treatments, the decrease was higher in heated than unheated communities indicating a lower resistance of heated communities to the drought. After only 13 days of recovery, green vegetation cover of both temperature treatments approached values similar to those observed before the imposed drought, suggesting similar resilience in both treatments. Above-ground biomass was reduced by elevated temperature, consistently in all S-levels, showing that the drought period did not change the biomass production patterns observed in the treatments before the imposed drought. Our results suggest that, regardless of the continuous exposure to elevated temperatures and associated short mild droughts, heated communities had not developed clear mechanisms to better cope with extended summer droughts.

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Abstract. Here we report on the single and combined impacts of climate warming and species richness on the biomass production in experimental grassland communities. Projections of a future warmer climate have stimulated studies on the response of terrestrial ecosystems to this global change. Experiments have likewise addressed the importance of species numbers for ecosystem functioning. There is, however, little knowledge on the interplay between warming and species richness. During three years, we grew experimental plant communities containing one, three or nine grassland species in 12 sunlit, climate-controlled chambers in Wilrijk, Belgium. Half of these chambers were exposed to ambient air temperatures (unheated), while the other half were warmed by 3 • C (heated). Equal amounts of water were added to heated and unheated communities, so that warming would imply drier soils if evapotranspiration was higher. Biomass production was decreased due to warming, both aboveground (-29%) and belowground (-25%), as negative impacts of increased heat and drought stress in summer prevailed. Complementarity effects, likely mostly through both increased aboveground spatial complementarity and facilitative effects of legumes, led to higher shoot and root biomass in multi-species communities, regardless of the induced warming. Surprisingly, warming suppressed productivity the most in 9-species communities, which may be attributed to negative impacts of intense interspecific competiCorrespondence to: H. J. De Boeck (hans.deboeck@ua.ac.be) tion for resources under conditions of high abiotic stress. Our results suggest that warming and the associated soil drying could reduce primary production in many temperate grasslands, and that this will not necessarily be mitigated by efforts to maintain or increase species richness.

Abstract.To study the effects of warming and declining species richness on the carbon balance of grassland communities, model ecosystems containing one, three or nine species were exposed to ambient and elevated (ambient +3 • C) air temperature. In this paper, we analyze measured ecosystem CO 2 fluxes to test whether ecosystem photosynthesis and respiration had acclimated to warming after 28 months of continuous heating, and whether the degree of acclimation depended on species richness. In order to test whether acclimation occurred, short term temperature response curves were established for all communities in both treatments. At similar temperatures, lower flux rates in the heated communities as compared to the unheated communities would indicate thermal acclimation. Because plant cover was significantly higher in the heated treatment, we normalized the data for plant cover. Subsequently, down-regulation of both photosynthesis and respiration was observed. Although CO 2 fluxes were larger in communities with higher species richness, species richness did not affect the degree of acclimation to warming. These results imply that models need to take thermal acclimation into account to simulate photosynthesis and respiration in a warmer world.