Functional trait expression of grassland species shift with short- and long-term nutrient additions

Humans are altering nutrient availability worldwide, likely affecting plant trait expression, with consequences for community composition and ecosystem function. Here, we examined the responses of plant species dominant under ambient nutrient conditions (baseline species) versus those that become dominant under increased nutrient conditions (enriched species) in a tallgrass prairie ecosystem. The expression of 8 functional traits was quantified for 3 baseline and 3 enriched species within one short-term and one long-term nutrient addition experiment. We found that enriched species occupied a trait space characterized by traits that generally correspond with faster growth rates than baseline species. Additionally, the enriched species shifted in their trait expression relative to the control more than the baseline species with nutrient additions, particularly within the long-term experiment. The trait space shifts of individual species with nutrient additions scaled up to affect community aggregate trait values within both experiments. However, traits that responded to nutrient additions at the community level were not strong predictors of aboveground net primary productivity (ANPP) within the short-term experiment. In contrast, in the long-term experiment, one response trait (community aggregate height) strongly correlated with variation in ANPP with nutrient additions. The link between plant functional traits and community and ecosystem responses to chronic nutrient additions shown here will provide important insight into key mechanisms driving grassland responses to global change.

[1]  Wilfried Thuiller,et al.  A multi‐trait approach reveals the structure and the relative importance of intra‐ vs. interspecific variability in plant traits , 2010 .

[2]  S. Polasky,et al.  Nutrient enrichment, biodiversity loss, and consequent declines in ecosystem productivity , 2013, Proceedings of the National Academy of Sciences.

[3]  P. Reich,et al.  A global study of relationships between leaf traits, climate and soil measures of nutrient fertility , 2009 .

[4]  Ulrike Groemping,et al.  Relative Importance for Linear Regression in R: The Package relaimpo , 2006 .

[5]  J. Funk.  Differences in plasticity between invasive and native plants from a low resource environment , 2008 .

[6]  Sandra Lavorel,et al.  How fundamental plant functional trait relationships scale‐up to trade‐offs and synergies in ecosystem services , 2012 .

[7]  Christina Gloeckner,et al.  Modern Applied Statistics With S , 2003 .

[8]  D. Tilman,et al.  Convergence and divergence of old-field vegetation after 11 yr of nitrogen addition , 1995 .

[9]  R. Cowling,et al.  Convergence in vegetation structure in the mediterranean communities of California, Chile and South Africa , 1980, Vegetatio.

[10]  J. Funk,et al.  A functional trait perspective on plant invasion. , 2012, Annals of botany.

[11]  W. Harpole,et al.  Nitrogen enrichment and plant communities , 2010, Annals of the New York Academy of Sciences.

[12]  W. K. Lauenroth,et al.  Inertia in Plant Community Structure: State Changes After Cessation of Nutrient‐Enrichment Stress , 1995 .

[13]  K. Gross,et al.  Functional- and abundance-based mechanisms explain diversity loss due to N fertilization. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[14]  David Tilman,et al.  Loss of plant species after chronic low-level nitrogen deposition to prairie grasslands , 2008, Nature.

[15]  Sandra Díaz,et al.  Scaling environmental change through the community‐level: a trait‐based response‐and‐effect framework for plants , 2008 .

[16]  Jennifer A Hoeting,et al.  A structured and dynamic framework to advance traits-based theory and prediction in ecology. , 2010, Ecology letters.

[17]  P. Reich,et al.  Strategy shifts in leaf physiology, structure and nutrient content between species of high‐ and low‐rainfall and high‐ and low‐nutrient habitats , 2001 .

[18]  Walter Krämer,et al.  Review of Modern applied statistics with S, 4th ed. by W.N. Venables and B.D. Ripley. Springer-Verlag 2002 , 2003 .

[19]  S. Prober,et al.  Plastic Traits of an Exotic Grass Contribute to Its Abundance but Are Not Always Favourable , 2012, PloS one.

[20]  David A. Wardle,et al.  Contrasting effects of plant inter‐ and intraspecific variation on community‐level trait measures along an environmental gradient , 2013 .

[21]  Meghan L. Avolio,et al.  Genetic diversity of a dominant C4 grass is altered with increased precipitation variability , 2012, Oecologia.

[22]  F. S. Chapin,et al.  The Mineral Nutrition of Wild Plants , 1980 .

[23]  Mark Westoby,et al.  Land-plant ecology on the basis of functional traits. , 2006, Trends in ecology & evolution.

[24]  W. Härdtle,et al.  Functional trait similarity of native and invasive herb species in subtropical China: environment-specific differences are the key , 2012 .

[25]  Olivier Honnay,et al.  A trait‐based analysis of the role of phosphorus vs. nitrogen enrichment in plant species loss across North‐west European grasslands , 2011 .

[26]  H. Mooney,et al.  Human Domination of Earth’s Ecosystems , 1997, Renewable Energy.

[27]  P. Reich,et al.  Assessing the generality of global leaf trait relationships. , 2005, The New phytologist.

[28]  R Core Team,et al.  R: A language and environment for statistical computing. , 2014 .

[29]  P. Reich,et al.  Leaf Life‐Span in Relation to Leaf, Plant, and Stand Characteristics among Diverse Ecosystems , 1992 .

[30]  F. Stuart Chapin,et al.  Evolution of Suites of Traits in Response to Environmental Stress , 1993, The American Naturalist.

[31]  Community-aggregated plant traits interact with soil nutrient heterogeneity to determine ecosystem functioning , 2013, Plant and Soil.

[32]  C. Violle,et al.  Let the concept of trait be functional , 2007 .

[33]  William N. Venables,et al.  Modern Applied Statistics with S , 2010 .

[34]  Jacob McC. Overton,et al.  Shifts in trait‐combinations along rainfall and phosphorus gradients , 2000 .

[35]  Helmut Hillebrand,et al.  Nutrient co-limitation of primary producer communities. , 2011, Ecology letters.

[36]  W. Stanley Harpole,et al.  Grassland species loss resulting from reduced niche dimension , 2007, Nature.

[37]  K. Thompson,et al.  The plant traits that drive ecosystems: Evidence from three continents , 2004 .

[38]  K. Treseder,et al.  Nitrogen limitation of net primary productivity in terrestrial ecosystems is globally distributed. , 2008, Ecology.

[39]  S. Díaz,et al.  Vive la différence: plant functional diversity matters to ecosystem processes , 2001 .

[40]  Yang Li,et al.  Plant community responses to nitrogen addition and increased precipitation: the importance of water availability and species traits , 2011 .

[41]  K. Reinhart,et al.  Specific Leaf Area Along a Nitrogen Fertilization Gradient , 2000 .

[42]  Brian J McGill,et al.  How do traits vary across ecological scales? A case for trait-based ecology. , 2010, Ecology letters.

[43]  J. Lepš,et al.  A test of the explanatory power of plant functional traits on the individual and population levels , 2011 .

[44]  P. Legendre,et al.  vegan : Community Ecology Package. R package version 1.8-5 , 2007 .

[45]  J. Fargione,et al.  Environmental and plant community determinants of species loss following nitrogen enrichment. , 2007, Ecology letters.

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

[47]  Stephen Porder,et al.  Terrestrial phosphorus limitation: mechanisms, implications, and nitrogen-phosphorus interactions. , 2010, Ecological applications : a publication of the Ecological Society of America.

[48]  J. P. Grime,et al.  Evidence for the Existence of Three Primary Strategies in Plants and Its Relevance to Ecological and Evolutionary Theory , 1977, The American Naturalist.

[49]  J. Schellberg,et al.  Plant functional traits and nutrient gradients on grassland , 2012 .

[50]  G. Likens,et al.  Technical Report: Human Alteration of the Global Nitrogen Cycle: Sources and Consequences , 1997 .

[51]  Sandra Lavorel,et al.  Using plant functional traits to understand the landscape distribution of multiple ecosystem services , 2011 .

[52]  Xu Yu-bo,et al.  Reachability Checking of Finite Precision Timed Automata , 2006 .

[53]  Katharine N. Suding,et al.  RELATIONSHIPS AMONG SPECIES TRAITS: SEPARATING LEVELS OF RESPONSE AND IDENTIFYING LINKAGES TO ABUNDANCE , 2003 .

[54]  A. Kinzig,et al.  Original Articles: Plant Attribute Diversity, Resilience, and Ecosystem Function: The Nature and Significance of Dominant and Minor Species , 1999, Ecosystems.

[55]  Ghislain Vieilledent,et al.  When and how should intraspecific variability be considered in trait-based plant ecology? , 2011 .

[56]  Katharine N Suding,et al.  Testing the Holy Grail framework: using functional traits to predict ecosystem change. , 2008, The New phytologist.

[57]  Vanessa Minden,et al.  Testing the effect–response framework: key response and effect traits determining above‐ground biomass of salt marshes , 2011 .

[58]  M. Westoby,et al.  Leaf-size divergence along rainfall and soil-nutrient gradients: Is the method of size reduction common among clades? , 2003 .

[59]  B. Enquist,et al.  Rebuilding community ecology from functional traits. , 2006, Trends in ecology & evolution.

[60]  Meghan L. Avolio,et al.  Changes in plant community composition, not diversity, during a decade of nitrogen and phosphorus additions drive above‐ground productivity in a tallgrass prairie , 2014 .

[61]  Hendrik Poorter,et al.  A comparison of specific leaf area, chemical composition and leaf construction costs of field plants from 15 habitats differing in productivity , 1999 .

[62]  Eric Garnier,et al.  PLANT FUNCTIONAL MARKERS CAPTURE ECOSYSTEM PROPERTIES DURING SECONDARY SUCCESSION , 2004 .

[63]  J. P. Grime,et al.  The plant traits that drive ecosystems: Evidence from three continents , 2004 .

[64]  D. Norton,et al.  Which plant traits determine abundance under long‐term shifts in soil resource availability and grazing intensity? , 2012 .

[65]  R. Pakeman,et al.  Multivariate identification of plant functional response and effect traits in an agricultural landscape. , 2011, Ecology.

[66]  Mark Westoby,et al.  EVOLUTIONARY DIVERGENCES IN LEAF STRUCTURE AND CHEMISTRY, COMPARING RAINFALL AND SOIL NUTRIENT GRADIENTS , 1999 .