Impacts of warming and elevated CO2 on a semi-arid grassland are non-additive, shift with precipitation, and reverse over time.

It is unclear how elevated CO2 (eCO2 ) and the corresponding shifts in temperature and precipitation will interact to impact ecosystems over time. During a 7-year experiment in a semi-arid grassland, the response of plant biomass to eCO2 and warming was largely regulated by interannual precipitation, while the response of plant community composition was more sensitive to experiment duration. The combined effects of eCO2 and warming on aboveground plant biomass were less positive in 'wet' growing seasons, but total plant biomass was consistently stimulated by ~ 25% due to unique, supra-additive responses of roots. Independent of precipitation, the combined effects of eCO2 and warming on C3 graminoids became increasingly positive and supra-additive over time, reversing an initial shift toward C4 grasses. Soil resources also responded dynamically and non-additively to eCO2 and warming, shaping the plant responses. Our results suggest grasslands are poised for drastic changes in function and highlight the need for long-term, factorial experiments.

[1]  Lauren M. Porensky,et al.  Thresholds and gradients in a semi‐arid grassland: long‐term grazing treatments induce slow, continuous and reversible vegetation change , 2016 .

[2]  J. Derner,et al.  Grazing intensity differentially regulates ANPP response to precipitation in North American semiarid grasslands. , 2016, Ecological applications : a publication of the Ecological Society of America.

[3]  S. Kim Authorship , 2016, Korean journal of family medicine.

[4]  Yiqi Luo,et al.  Evidence for long‐term shift in plant community composition under decadal experimental warming , 2015 .

[5]  Rhonda Hoenigman,et al.  Seasonality of soil moisture mediates responses of ecosystem phenology to elevated CO2 and warming in a semi‐arid grassland , 2015 .

[6]  E. Pendall,et al.  Antecedent moisture and temperature conditions modulate the response of ecosystem respiration to elevated CO2 and warming , 2015, Global change biology.

[7]  A. Dai,et al.  The Magnitude and Causes of Global Drought Changes in the Twenty-First Century under a Low-Moderate Emissions Scenario , 2015 .

[8]  Atul K. Jain,et al.  The dominant role of semi-arid ecosystems in the trend and variability of the land CO2 sink , 2015, Science.

[9]  S. Frey,et al.  Global environmental change and the nature of aboveground net primary productivity responses: insights from long-term experiments , 2015, Oecologia.

[10]  E. Pendall,et al.  Microclimatic Performance of a Free-Air Warming and CO2 Enrichment Experiment in Windy Wyoming, USA , 2015, PloS one.

[11]  E. Pendall,et al.  Long-term exposure to elevated CO2 enhances plant community stability by suppressing dominant plant species in a mixed-grass prairie , 2014, Proceedings of the National Academy of Sciences.

[12]  P. Newton,et al.  Seasonal not annual rainfall determines grassland biomass response to carbon dioxide , 2014, Nature.

[13]  Yi Y. Liu,et al.  Contribution of semi-arid ecosystems to interannual variability of the global carbon cycle , 2014, Nature.

[14]  Gregory S. McMaster,et al.  Elevated CO2 further lengthens growing season under warming conditions , 2014, Nature.

[15]  F. Woodward,et al.  Physiological advantages of C4 grasses in the field: a comparative experiment demonstrating the importance of drought , 2014, Global change biology.

[16]  Y. Carrillo,et al.  Disentangling root responses to climate change in a semiarid grassland , 2014, Oecologia.

[17]  M. Lieffering,et al.  Selective grazing modifies previously anticipated responses of plant community composition to elevated CO2 in a temperate grassland , 2014, Global change biology.

[18]  Y. Carrillo,et al.  Warming Reduces Carbon Losses from Grassland Exposed to Elevated Atmospheric Carbon Dioxide , 2013, PloS one.

[19]  Laura Turnbull,et al.  Sensitivity of grassland plant community composition to spatial vs. temporal variation in precipitation. , 2013, Ecology.

[20]  P. Reich,et al.  Effects of plant diversity, N fertilization, and elevated carbon dioxide on grassland soil N cycling in a long‐term experiment , 2013, Global change biology.

[21]  P. Reich,et al.  Decade-long soil nitrogen constraint on the CO2 fertilization of plant biomass , 2013 .

[22]  F. Dijkstra,et al.  Climate change reduces the net sink of CH4 and N2O in a semiarid grassland , 2012, Global change biology.

[23]  H. Tian,et al.  Effects of Elevated Carbon Dioxide and Increased Temperature on Methane and Nitrous Oxide Fluxes: Evidence from Field Experiments , 2012 .

[24]  O. Sala,et al.  Legacies of precipitation fluctuations on primary production: theory and data synthesis , 2012, Philosophical Transactions of the Royal Society B: Biological Sciences.

[25]  Y. Carrillo,et al.  Climate change alters stoichiometry of phosphorus and nitrogen in a semiarid grassland. , 2012, The New phytologist.

[26]  V. Jin,et al.  Feedback from plant species change amplifies CO2 enhancement of grassland productivity , 2012, Global change biology.

[27]  F. Dijkstra,et al.  Simple additive effects are rare: a quantitative review of plant biomass and soil process responses to combined manipulations of CO2 and temperature , 2012, Global change biology.

[28]  B. Hungate,et al.  Biogeochemical and ecological feedbacks in grassland responses to warming , 2012 .

[29]  Y. Carrillo,et al.  Controls over Soil Nitrogen Pools in a Semiarid Grassland Under Elevated CO2 and Warming , 2012, Ecosystems.

[30]  Jizhong Zhou,et al.  Interannual variability in responses of belowground net primary productivity (NPP) and NPP partitioning to long‐term warming and clipping in a tallgrass prairie , 2012 .

[31]  S. Hoeppner,et al.  Interactive responses of old‐field plant growth and composition to warming and precipitation , 2012 .

[32]  S. Heckathorn,et al.  A meta-analysis of plant physiological and growth responses to temperature and elevated CO2 , 2012, Oecologia.

[33]  A. Dai Drought under global warming: a review , 2011 .

[34]  Alan K. Knapp,et al.  Relative effects of precipitation variability and warming on tallgrass prairie ecosystem function , 2011 .

[35]  Mark West,et al.  C4 grasses prosper as carbon dioxide eliminates desiccation in warmed semi-arid grassland , 2011, Nature.

[36]  T. Barrett Computations using analysis of covariance , 2011 .

[37]  J. Morgan,et al.  Application of a conceptual framework to interpret variability in rangeland responses to atmospheric CO2 enrichment , 2010, The Journal of Agricultural Science.

[38]  F. Dijkstra,et al.  Elevated CO2 effects on semi-arid grassland plants in relation to water availability and competition. , 2010 .

[39]  M. Lieffering,et al.  The rate of progression and stability of progressive nitrogen limitation at elevated atmospheric CO2 in a grazed grassland over 11 years of Free Air CO2 enrichment , 2010, Plant and Soil.

[40]  Y. Carrillo,et al.  Contrasting effects of elevated CO2 and warming on nitrogen cycling in a semiarid grassland. , 2010, The New phytologist.

[41]  D. Taub,et al.  Interactive effects of elevated carbon dioxide and environmental stresses on root mass fraction in plants: a meta-analytical synthesis using pairwise techniques , 2010, Oecologia.

[42]  F. Woodward,et al.  Ecophysiological traits in C3 and C4 grasses: a phylogenetically controlled screening experiment. , 2010, The New phytologist.

[43]  S. Collins,et al.  A framework for assessing ecosystem dynamics in response to chronic resource alterations induced by global change. , 2009, Ecology.

[44]  A. Rogers,et al.  Elevated CO2 effects on plant carbon, nitrogen, and water relations: six important lessons from FACE. , 2009, Journal of experimental botany.

[45]  K. Havstad,et al.  Grazing Intensity on Vegetation Dynamics of a Typical Steppe in Northeast Inner Mongolia , 2009 .

[46]  P. Ciais,et al.  Modeled interactive effects of precipitation, temperature, and [CO2] on ecosystem carbon and water dynamics in different climatic zones , 2008 .

[47]  Yiqi Luo Terrestrial Carbon-Cycle Feedback to Climate Warming , 2007 .

[48]  Mark West,et al.  Carbon dioxide enrichment alters plant community structure and accelerates shrub growth in the shortgrass steppe , 2007, Proceedings of the National Academy of Sciences.

[49]  W. Parton,et al.  Projected ecosystem impact of the Prairie Heating and CO2 Enrichment experiment. , 2007, The New phytologist.

[50]  J. Derner,et al.  Grazing-Induced Modifications to Peak Standing Crop in Northern Mixed-grass Prairie , 2007 .

[51]  A. Rogers,et al.  The response of photosynthesis and stomatal conductance to rising [CO2]: mechanisms and environmental interactions. , 2007, Plant, cell & environment.

[52]  M. Power,et al.  Species Interactions Reverse Grassland Responses to Changing Climate , 2007, Science.

[53]  P. Reich,et al.  Carbon-Nitrogen Interactions in Terrestrial Ecosystems in Response to Rising Atmospheric Carbon Dioxide , 2006 .

[54]  Zhen Liu,et al.  The response of a shrub‐invaded grassland on the Inner Mongolia steppe to long‐term grazing by sheep , 2006 .

[55]  A. Prokushkin,et al.  Critical analysis of root : shoot ratios in terrestrial biomes , 2006 .

[56]  P. Reich,et al.  Linking leaf and root trait syndromes among 39 grassland and savannah species. , 2005, The New phytologist.

[57]  P. Nyren,et al.  Competition between Artemisia frigida and Cleistogenes squarrosa under different clipping intensities in replacement series mixtures at different nitrogen levels , 2005 .

[58]  W. Parton,et al.  Progressive Nitrogen Limitation of Ecosystem Responses to Rising Atmospheric Carbon Dioxide , 2004 .

[59]  M. R. Shaw,et al.  Water relations in grassland and desert ecosystems exposed to elevated atmospheric CO2 , 2004, Oecologia.

[60]  R. Norby,et al.  Evaluating ecosystem responses to rising atmospheric CO2 and global warming in a multi‐factor world , 2004 .

[61]  D. Ellsworth,et al.  Functional responses of plants to elevated atmospheric CO2– do photosynthetic and productivity data from FACE experiments support early predictions? , 2004 .

[62]  F. Samson,et al.  Great Plains ecosystems: past, present, and future , 2004 .

[63]  J. C. Winslow,et al.  The influence of seasonal water availability on global C3 versus C4 grassland biomass and its implications for climate change research , 2003 .

[64]  P. Reich,et al.  Functional traits, productivity and effects on nitrogen cycling of 33 grassland species , 2002 .

[65]  Kenneth A. Lachlan,et al.  The high cost of complexity in experimental design and data analysis: Type I and type II error rates in multiway ANOVA , 2002 .

[66]  P. Reich,et al.  Do species and functional groups differ in acquisition and use of C, N and water under varying atmospheric CO2 and N availability regimes? : a field test with 16 grassland species , 2001 .

[67]  A. Knapp,et al.  Variation among biomes in temporal dynamics of aboveground primary production. , 2001, Science.

[68]  F. Chapin,et al.  Global Warming and Terrestrial Ecosystems: A Conceptual Framework for Analysis , 2000 .

[69]  Richard H. Hart,et al.  IMPACT OF GRAZING MANAGEMENT ON THE CARBON AND NITROGEN BALANCE OF A MIXED-GRASS RANGELAND , 1999 .

[70]  P. Martikainen,et al.  Global change and root function , 1998 .

[71]  Howard E. Epstein,et al.  Plant Effects on Spatial and Temporal Patterns of Nitrogen Cycling in Shortgrass Steppe , 1998, Ecosystems.

[72]  D. Tilman,et al.  Influence of Nitrogen Loading and Species Composition on the Carbon Balance of Grasslands , 1996, Science.

[73]  K. Kindscher,et al.  Prairie plant guilds: a multivariate analysis of prairie species based on ecological and morphological traits , 1995, Vegetatio.

[74]  S. Stehman,et al.  Practical analysis of factorial experiments in forestry , 1995 .

[75]  S. Wilson,et al.  Competition, Fire, and Nutients in a Mixed-Grass Prairie , 1990 .

[76]  J. L. Dodd,et al.  Phenological Pattern in the Shortgrass Prairie , 1976 .

[77]  P. Jones,et al.  Global warming and changes in drought , 2014 .

[78]  J. Liebmann,et al.  Rate of Progression , 2014 .

[79]  G. Yohe,et al.  Climate Change Impacts in the United States: The Third National Climate Assessment , 2014 .

[80]  P. Reich,et al.  Biomass allocation to leaves, stems and roots: meta-analyses of interspecific variation and environmental control. , 2012, The New phytologist.

[81]  R. Sage,et al.  Quo vadis C4? An ecophysiological perspective on global change and the future of C4 plants , 2004, Photosynthesis Research.

[82]  W. K. Lauenroth,et al.  The effects of water- and nitrogen-induced stresses on plant community structure in a semiarid grassland , 2004, Oecologia.

[83]  G. Daily,et al.  Ecosystem services in grasslands. , 1997 .