title: Biotic context and soil properties modulate native plant responses to enhanced rainfall.
abstract: OBJECTIVE
  The environmental and biotic context within which plants grow have a great potential to modify responses to climatic changes, yet few studies have addressed both the direct effects of climate and the modulating roles played by variation in the biotic (e.g. competitors) and abiotic (e.g. soils) environment.
  
  
  METHODS
  In a grassland with highly heterogeneous soils and community composition, small seedlings of two native plants, Lasthenia californica and Calycadenia pauciflora, were transplanted into factorially watered and fertilized plots. Measurements were made to test how the effect of climatic variability (mimicked by the watering treatment) on the survival, growth and seed production of these species was modulated by above-ground competition and by edaphic variables.
  
  
  RESULTS
  Increased competition outweighed the direct positive impacts of enhanced rainfall on most fitness measures for both species, resulting in no net effect of enhanced rainfall. Both species benefitted from enhanced rainfall when the absence of competitors was accompanied by high soil water retention capacity. Fertilization did not amplify the watering effects; rather, plants benefitted from enhanced rainfall or competitor removal only in ambient nutrient conditions with high soil water retention capacity.
  
  
  CONCLUSIONS
  The findings show that the direct effects of climatic variability on plant fitness may be reversed or neutralized by competition and, in addition, may be strongly modulated by soil variation. Specifically, coarse soil texture was identified as a factor that may limit plant responsiveness to altered water availability. These results highlight the importance of considering the abiotic as well as biotic context when making future climate change forecasts.
authors.affiliation: Department of Environmental Science and Policy, University of California Davis, One Shields Avenue, Davis, CA 95616, USA and Department of Biology, University of Oulu, PO Box 3000, FI-90014 University of Oulu, Finland anu.eskelinen@oulu.fi.
authors.forename: Anu
authors.initials: A
authors.lastname: Eskelinen
authors.affiliation: Department of Environmental Science and Policy, University of California Davis, One Shields Avenue, Davis, CA 95616, USA and.
authors.forename: Susan
authors.initials: S
authors.lastname: Harrison
mesh_terms: D019659:Asteraceae / Q000254:growth & development / Q000502:physiology*; D018533:Biomass; D002980:Climate; D057231:Climate Change; D065948:Grassland; D016014:Linear Models; D011891:Rain; D012987:Soil / Q000737:chemistry*; D014867:Water / Q000502:physiology*
chemical_list: D012987:Soil; D014867:Water
keywords: Biotic context; California annual natives; Calycadenia pauciflora; Lasthenia californica; climate change; competition; direct and indirect effects; enhanced rainfall; floral herbivory; grassland; multiple global changes; plant–climate interactions; soil properties