Scaling environmental change through the community‐level: a trait‐based response‐and‐effect framework for plants

Predicting ecosystem responses to global change is a major challenge in ecology. A critical step in that challenge is to understand how changing environmental conditions influence processes across levels of ecological organization. While direct scaling from individual to ecosystem dynamics can lead to robust and mechanistic predictions, new approaches are needed to appropriately translate questions through the community level. Species invasion, loss, and turnover all necessitate this scaling through community processes, but predicting how such changes may influence ecosystem function is notoriously difficult. We suggest that community-level dynamics can be incorporated into scaling predictions using a trait-based response-effect framework that differentiates the community response to environmental change (predicted by response traits) and the effect of that change on ecosystem processes (predicted by effect traits). We develop a response-and-effect functional framework, concentrating on how the relationships among species' response, effect, and abundance can lead to general predictions concerning the magnitude and direction of the influence of environmental change on function. We then detail several key research directions needed to better scale the effects of environmental change through the community level. These include (1) effect and response trait characterization, (2) linkages between response-and-effect traits, (3) the importance of species interactions on trait expression, and (4) incorporation of feedbacks across multiple temporal scales. Increasing rates of extinction and invasion that are modifying communities worldwide make such a research agenda imperative.

[1]  N. Williams,et al.  Extinction order and altered community structure rapidly disrupt ecosystem functioning. , 2005, Ecology letters.

[2]  F. Keesing,et al.  Frontiers of Ecology , 2001 .

[3]  N. LeRoy Poff,et al.  Landscape Filters and Species Traits: Towards Mechanistic Understanding and Prediction in Stream Ecology , 1997, Journal of the North American Benthological Society.

[4]  M. Rejmánek,et al.  Toward a Causal Explanation of Plant Invasiveness: Seedling Growth and Life‐History Strategies of 29 Pine (Pinus) Species , 2002, The American Naturalist.

[5]  J. Castilla,et al.  Challenges in the Quest for Keystones , 1996 .

[6]  C. D’Antonio,et al.  COMPETITION BETWEEN NATIVE PERENNIAL AND EXOTIC ANNUAL GRASSES: IMPLICATIONS FOR AN HISTORICAL INVASION , 2004 .

[7]  D. Ackerly,et al.  Plant growth and reproduction along CO2 gradients: non‐linear responses and implications for community change , 1995 .

[8]  J. Overpeck,et al.  Responses of plant populations and communities to environmental changes of the late Quaternary , 2000, Paleobiology.

[9]  Mark W. Schwartz,et al.  Rare Species and Ecosystem Functioning , 2005 .

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

[11]  D. Goldberg,et al.  Equivalence of competitors in plant communities: a null hypothesis and a field experimental approach. , 1983 .

[12]  Steven I. Higgins,et al.  Estimating plant migration rates under habitat loss and fragmentation , 2003 .

[13]  S. Carpenter,et al.  Ecological forecasts: an emerging imperative. , 2001, Science.

[14]  Owen L. Petchey,et al.  Extinction and the loss of functional diversity , 2002, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[15]  M. Cadotte,et al.  Life‐history correlates of plant invasiveness at regional and continental scales , 2005 .

[16]  S. Díaz,et al.  Can grazing response of herbaceous plants be predicted from simple vegetative traits , 2001 .

[17]  Mark Rees,et al.  Decoupling the direct and indirect effects of nitrogen deposition on ecosystem function. , 2006, Ecology letters.

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

[19]  J. Bruno,et al.  Inclusion of facilitation into ecological theory , 2003 .

[20]  Marten Scheffer,et al.  Large Species Shifts Triggered by Small Forces , 2004, The American Naturalist.

[21]  A. Prasad,et al.  Potential Changes in Tree Species Richness and Forest Community Types following Climate Change , 2001, Ecosystems.

[22]  James B. Grace,et al.  Components of resource competition in plant communities. , 1990 .

[23]  Chris Margules,et al.  Which traits of species predict population declines in experimental forest fragments , 2000 .

[24]  S. Lavorel,et al.  Mechanisms underlying the impacts of exotic plant invasions , 2003, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[25]  T. Swetnam,et al.  Mesoscale Disturbance and Ecological Response to Decadal Climatic Variability in the American Southwest , 1998 .

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

[27]  Kate E. Jones,et al.  Multiple Causes of High Extinction Risk in Large Mammal Species , 2005, Science.

[28]  D. Wardle,et al.  The response of a three trophic level soil food web to the identity and diversity of plant species and functional groups , 2003 .

[29]  Cynthia S. Brown,et al.  Community assembly and invasion: An experimental test of neutral versus niche processes , 2003, Proceedings of the National Academy of Sciences of the United States of America.

[30]  D. Wardle,et al.  Ecological Linkages Between Aboveground and Belowground Biota , 2004, Science.

[31]  Paul A. Keddy,et al.  Community Assembly Rules, Morphological Dispersion, and the Coexistence of Plant Species , 1998 .

[32]  M. Rietkerk,et al.  The Dynamic Regime Concept for Ecosystem Management and Restoration , 2004 .

[33]  F. Chapin,et al.  Plant and soil responses to neighbour removal and fertilization in Alaskan tussock tundra , 2004 .

[34]  P. Vitousek,et al.  Biological invasions by exotic grasses, the grass/fire cycle, and global change , 1992 .

[35]  D. Mouillot,et al.  Niche overlap estimates based on quantitative functional traits: a new family of non-parametric indices , 2005, Oecologia.

[36]  S. Hobbie Effects of plant species on nutrient cycling. , 1992, Trends in ecology & evolution.

[37]  J. Wootton Mechanisms of successional dynamics: Consumers and the rise and fall of species dominance , 2002, Ecological Research.

[38]  C. D’Antonio,et al.  Variation in the impact of exotic grasses on native plant composition in relation to fire across an elevation gradient in Hawaii , 2000 .

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

[40]  Shawn J. Marshall,et al.  Freshwater Forcing of Abrupt Climate Change During the Last Glaciation , 2001, Science.

[41]  O. Eriksson,et al.  Litter species composition influences the performance of seedlings of grassland herbs , 2006 .

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

[43]  F. Stuart Chapin,et al.  THE RESPONSE OF TUNDRA PLANT BIOMASS, ABOVEGROUND PRODUCTION, NITROGEN, AND CO2 FLUX TO EXPERIMENTAL WARMING , 1998 .

[44]  T. Miller,et al.  Dispersal Rates Affect Species Composition in Metacommunities of Sarracenia purpurea Inquilines , 2003, The American Naturalist.

[45]  J. Kutzbach,et al.  A severe centennial-scale drought in midcontinental North America 4200 years ago and apparent global linkages , 2005 .

[46]  Daniel G. Milchunas,et al.  Plant trait responses to grazing – a global synthesis , 2007 .

[47]  Michael A. McCarthy,et al.  Plant traits and local extinctions in natural grasslands along an urban–rural gradient , 2005 .

[48]  Dylan G. Fischer,et al.  COMMUNITY AND ECOSYSTEM GENETICS: A CONSEQUENCE OF THE EXTENDED PHENOTYPE , 2003 .

[49]  D. Wardle,et al.  Interspecific interactions and biomass allocation among grassland plant species , 2003 .

[50]  Robert K. Colwell,et al.  Species Loss and Aboveground Carbon Storage in a Tropical Forest , 2005, Science.

[51]  P. Reich,et al.  The Evolution of Plant Functional Variation: Traits, Spectra, and Strategies , 2003, International Journal of Plant Sciences.

[52]  J. Cavender-Bares,et al.  Phylogenetic Overdispersion in Floridian Oak Communities , 2004, The American Naturalist.

[53]  D. Goldberg Competitive ability: Definitions, contingency and correlated traits , 1996 .

[54]  James S. Clark,et al.  STAGES AND SPATIAL SCALES OF RECRUITMENT LIMITATION IN SOUTHERN APPALACHIAN FORESTS , 1998 .

[55]  O. Sala,et al.  Long-Term Forage Production of North American Shortgrass Steppe. , 1992, Ecological applications : a publication of the Ecological Society of America.

[56]  M. Bertness,et al.  LATITUDINAL DIFFERENCES IN PLANT PALATABILITY IN ATLANTIC COAST SALT MARSHES , 2001 .

[57]  K. Engelhardt Relating effect and response traits in submersed aquatic macrophytes. , 2006, Ecological applications : a publication of the Ecological Society of America.

[58]  F. Bello,et al.  Predictive value of plant traits to grazing along a climatic gradient in the Mediterranean , 2005 .

[59]  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 .

[60]  Harold A. Mooney,et al.  Effects of rainfall variability and gopher disturbance on serpentine annual grassland dynamics , 1991 .

[61]  S. Lavorel,et al.  Plant functional classifications: from general groups to specific groups based on response to disturbance. , 1997, Trends in ecology & evolution.

[62]  S. Naeem,et al.  Disentangling biodiversity effects on ecosystem functioning: deriving solutions to a seemingly insurmountable problem , 2003 .

[63]  F. Chapin,et al.  Non‐equilibrium succession dynamics indicate continued northern migration of lodgepole pine , 2003 .

[64]  R. O'Neill,et al.  The value of the world's ecosystem services and natural capital , 1997, Nature.

[65]  J. P. Grime,et al.  The response of two contrasting limestone grasslands to simulated climate change. , 2000, Science.

[66]  C. Lortie,et al.  Positive interactions among alpine plants increase with stress , 2002, Nature.

[67]  J. Lancaster,et al.  Assembly rules within a contingent ecology , 1999 .

[68]  J. L. Gittleman,et al.  Predicting extinction risk in declining species , 2000, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[69]  Sean C. Thomas,et al.  The worldwide leaf economics spectrum , 2004, Nature.

[70]  F. Woodward,et al.  Effects of elevated concentrations of carbon dioxide on individual plants, populations, communities and ecosystems , 1991 .

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

[72]  J. Betancourt,et al.  Role of multidecadal climate variability in a range extension of pinyon pine. , 2006, Ecology.

[73]  J. Arendt,et al.  Adaptive Intrinsic Growth Rates: An Integration Across Taxa , 1997, The Quarterly Review of Biology.

[74]  C. Daehler,et al.  Experimental Restoration of an Indigenous Hawaiian Grassland after Invasion by Buffel Grass (Cenchrus ciliaris) , 2005 .

[75]  José M. V. Fragoso,et al.  Forecasting Regional to Global Plant Migration in Response to Climate Change , 2005 .

[76]  DEBRA P. C. PETERS,et al.  Long-Term and Large-Scale Perspectives on the Relationship between Biodiversity and Ecosystem Functioning , 2003 .

[77]  Monica G. Turner,et al.  Filling key gaps in population and community ecology , 2007 .

[78]  Monica G. Turner,et al.  Ecological Thresholds: The Key to Successful Environmental Management or an Important Concept with No Practical Application? , 2006, Ecosystems.

[79]  F. Chapin,et al.  Consequences of changing biodiversity , 2000, Nature.

[80]  S. Hubbell,et al.  The unified neutral theory of biodiversity and biogeography at age ten. , 2011, Trends in ecology & evolution.

[81]  W. Silver,et al.  Effects of Global Changes on Above- and Belowground Biodiversity in Terrestrial Ecosystems: Implications for Ecosystem Functioning , 2000 .

[82]  Clive G. Jones,et al.  PREDICTING EFFECTS OF ECOSYSTEM ENGINEERS ON PATCH‐SCALE SPECIES RICHNESS FROM PRIMARY PRODUCTIVITY , 2004 .

[83]  P. Reich,et al.  Legume presence increases photosynthesis and N concentrations of co-occurring non-fixers but does not modulate their responsiveness to carbon dioxide enrichment , 2003, Oecologia.

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

[85]  T. Sharkey,et al.  Stomatal conductance and photosynthesis , 1982 .

[86]  O. Schmitz Perturbation and abrupt shift in trophic control of biodiversity and productivity , 2004 .

[87]  F. Bazzaz 14 – Scaling in Biological Systems: Population and Community Perspectives , 1993 .

[88]  J. P. Grime,et al.  Benefits of plant diversity to ecosystems: immediate, filter and founder effects , 1998 .

[89]  Sandra Lavorel,et al.  Grazing response groups among understorey plants in arid rangelands , 1999 .

[90]  Lydia Olander,et al.  Nitrogen and Nature , 2002, Ambio.

[91]  S. Peacor,et al.  The growth–mortality tradeoff: evidence from anuran larvae and consequences for species distributions , 2006, Oecologia.

[92]  D. Ackerly,et al.  A trait-based test for habitat filtering: convex hull volume. , 2006, Ecology.

[93]  David J. Currie,et al.  Projected Effects of Climate Change on Patterns of Vertebrate and Tree Species Richness in the Conterminous United States , 2001, Ecosystems.

[94]  U. Schurr,et al.  Changing the way we think about global change research: scaling up in experimental ecosystem science , 2004 .

[95]  F. Woodward,et al.  Functional Approaches to Predicting the Ecological Effects of Global Change , 1991 .

[96]  W. Reid,et al.  Millennium Ecosystem Assessment , 2005 .

[97]  A. L. Buikema,et al.  Do similar communities develop in similar sites? A test with zooplankton structure and function , 1998 .

[98]  W. Resetarits,et al.  Predator identity and ecological impacts: Functional redundancy or functional diversity? , 2003 .

[99]  Owen L. Petchey,et al.  Functional diversity: back to basics and looking forward. , 2006, Ecology letters.

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

[101]  Christian Körner,et al.  Biosphere responses to CO2 enrichment. , 2000 .

[102]  F. Stuart Chapin,et al.  Detecting changes in arctic tundra plant communities in response to warming over decadal time scales , 2004 .

[103]  Clarence Lehman,et al.  Conventional functional classification schemes underestimate the relationship with ecosystem functioning. , 2006, Ecology letters.

[104]  Roderick Hunt,et al.  Comparative Plant Ecology: A Functional Approach to Common British Species , 1989 .

[105]  Amy J. Symstad,et al.  Species diversity, functional diversity, and ecosystem functioning , 2002 .

[106]  Martin Solan,et al.  Extinction and Ecosystem Function in the Marine Benthos , 2004, Science.

[107]  F. Pugnaire,et al.  Changes in plant interactions along a gradient of environmental stress , 2001 .

[108]  D. Read,et al.  Specialized cheating of the ectomycorrhizal symbiosis by an epiparasitic liverwort , 2003, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[109]  W. Steffen,et al.  Global Change and the Earth System: A Planet Under Pressure , 2005 .

[110]  Garry D. Peterson,et al.  Response diversity, ecosystem change, and resilience , 2003 .

[111]  D. Tilman,et al.  QUADRATIC VARIATION IN OLD-FIELD SPECIES RICHNESS ALONG GRADIENTS OF DISTURBANCE AND NITROGEN , 2002 .

[112]  D. Wardle,et al.  Herbivore-mediated linkages between aboveground and belowground communities , 2003 .

[113]  F. Woodward Predicting plant responses to global environmental change. , 1992, The New phytologist.

[114]  J. P. Grime,et al.  Trait convergence and trait divergence in herbaceous plant communities: Mechanisms and consequences , 2006 .

[115]  Ricard V. Solé,et al.  Competition and introduction regime shape exotic bird communities in Hawaii , 2005, Biological Invasions.

[116]  J. P. Grime,et al.  Interacting effects of herbivory and fertility on a synthesized plant community , 1999 .

[117]  O. Phillips,et al.  Extinction risk from climate change , 2004, Nature.

[118]  Harold A. Mooney,et al.  Community and population dynamics of serpentine grassland annuals in relation to gopher disturbance , 1985, Oecologia.

[119]  Nitrogen limitation in dryland ecosystems: Responses to geographical and temporal variation in precipitation , 1999 .

[120]  Martin J. Wassen,et al.  Trophic interactions in a changing world , 2004 .

[121]  A. Knapp,et al.  Dominance not richness determines invasibility of tallgrass prairie. , 2004 .

[122]  D. Tilman,et al.  PLANT COMPETITION AND RESOURCE AVAILABILITY IN RESPONSE TO DISTURBANCE AND FERTILIZATION , 1993 .

[123]  S. Lavorel,et al.  Plant life-history attributes: their relationship to disturbance response in herbaceous vegetation. , 1995 .

[124]  M. Roderick,et al.  Challenging Theophrastus: A common core list of plant traits for functional ecology , 1999 .

[125]  A. Agrawal,et al.  Direct and interactive effects of enemies and mutualists on plant performance: a meta-analysis. , 2007, Ecology.

[126]  K. Klanderud Climate change effects on species interactions in an alpine plant community , 2005 .

[127]  S. Lavorel,et al.  Predicting changes in community composition and ecosystem functioning from plant traits: revisiting the Holy Grail , 2002 .

[128]  Robert Costanza,et al.  Economic Reasons for Conserving Wild Nature , 2002, Science.

[129]  O. Schmitz,et al.  Ecosystem Responses to Global Climate Change: Moving Beyond Color Mapping , 2003 .

[130]  M. Oesterheld,et al.  Effect of defoliation intensity on aboveground and belowground relative growth rates , 1992, Oecologia.

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

[132]  M. Willig,et al.  The Relationship Between Productivity and Species Richness , 1999 .

[133]  D. Spittlehouse,et al.  GENETIC RESPONSES TO CLIMATE IN PINUS CONTORTA: NICHE BREADTH, CLIMATE CHANGE, AND REFORESTATION , 1999 .

[134]  D. Tilman Resource competition and community structure. , 1983, Monographs in population biology.

[135]  P. Reich,et al.  From tropics to tundra: global convergence in plant functioning. , 1997, Proceedings of the National Academy of Sciences of the United States of America.

[136]  S. Lavorel,et al.  PLANT TRAITS IN A STATE AND TRANSITION FRAMEWORK AS MARKERS OF ECOSYSTEM RESPONSE TO LAND-USE CHANGE , 2007 .

[137]  F. Stuart Chapin,et al.  Plant functional types as predictors of transient responses of arctic vegetation to global change , 1996 .

[138]  Miguel Martínez-Ramos,et al.  APPLYING COMMUNITY STRUCTURE ANALYSIS TO ECOSYSTEM FUNCTION: EXAMPLES FROM POLLINATION AND CARBON STORAGE , 2005 .

[139]  C. Körner,et al.  A field study of the effects of elevated CO2 on plant biomass and community structure in a calcareous grassland , 1999, Oecologia.

[140]  M. Navas,et al.  Plant growth and competition at elevated CO2 : on winners, losers and functional groups. , 2003, The New phytologist.

[141]  Julio L. Betancourt,et al.  INFLUENCE OF LANDSCAPE STRUCTURE AND CLIMATE VARIABILITY ON A LATE HOLOCENE PLANT MIGRATION , 2003 .

[142]  Amy J. Symstad,et al.  Functional diversity revealed by removal experiments , 2003 .

[143]  S. Díaz,et al.  Plant functional types and ecosystem function in relation to global change , 1997 .

[144]  D. Wardle,et al.  Determinants of litter mixing effects in a Swedish boreal forest , 2003 .

[145]  C. Field,et al.  Interactive effects of elevated CO2, N deposition and climate change on plant litter quality in a California annual grassland , 2004, Oecologia.

[146]  F. Chapin,et al.  Species‐specific responses of plant communities to altered carbon and nutrient availability , 2001 .

[147]  John E. Kutzbach,et al.  Projected distributions of novel and disappearing climates by 2100 AD , 2006, Proceedings of the National Academy of Sciences.

[148]  M. Nyström Redundancy and Response Diversity of Functional Groups: Implications for the Resilience of Coral Reefs , 2006, Ambio.

[149]  N. LeRoy Poff,et al.  Life-history strategies predict fish invasions and extirpations in the colorado river basin , 2006 .

[150]  Craig R. Allen,et al.  Functional Group Change within and across Scales following Invasions and Extinctions in the Everglades Ecosystem , 2002, Ecosystems.

[151]  Sandra Díaz Elevated CO 2 Responsiveness, Interactions at the Community Level and Plant Functional Types , 1995 .

[152]  C. Kremen Managing ecosystem services: what do we need to know about their ecology? , 2005, Ecology letters.

[153]  Ian R. Noble,et al.  A functional classification for predicting the dynamics of landscapes , 1996 .

[154]  E. Schulze,et al.  Relationships among Maximum Stomatal Conductance, Ecosystem Surface Conductance, Carbon Assimilation Rate, and Plant Nitrogen Nutrition: A Global Ecology Scaling Exercise , 1994 .

[155]  R. May,et al.  Predicted correspondence between species abundances and dendrograms of niche similarities , 2003, Proceedings of the National Academy of Sciences of the United States of America.

[156]  F. Chapin,et al.  EFFECTS OF BIODIVERSITY ON ECOSYSTEM FUNCTIONING: A CONSENSUS OF CURRENT KNOWLEDGE , 2005 .

[157]  R. Marquis,et al.  ECOSYSTEM ENGINEERING BY CATERPILLARS INCREASES INSECT HERBIVORE DIVERSITY ON WHITE OAK , 2003 .

[158]  D. Ackerly Community Assembly, Niche Conservatism, and Adaptive Evolution in Changing Environments , 2003, International Journal of Plant Sciences.

[159]  S. Kark,et al.  Impacts and extent of biotic invasions in terrestrial ecosystems. , 2002 .

[160]  Christopher B. Field,et al.  RESPONSES OF TERRESTRIAL ECOSYSTEMS TO THE CHANGING ATMOSPHERE: A Resource-Based Approach*'** , 1992 .

[161]  J. P. Grime,et al.  The impact of elevated CO2 on plant-herbivore interactions: experimental evidence of moderating effects at the community level , 1998, Oecologia.

[162]  S. Allen,et al.  Aspects of the mineral nutrition of some native British plants — inter-site variation , 1987, Vegetatio.

[163]  D. Ackerly,et al.  A trait-based approach to community assembly: partitioning of species trait values into within- and among-community components. , 2007, Ecology letters.

[164]  J. Lambrinos THE VARIABLE INVASIVE SUCCESS OF CORTADERIA SPECIES IN A COMPLEX LANDSCAPE , 2002 .

[165]  O. Dangles,et al.  Simulating species loss following perturbation: assessing the effects on process rates , 2002, Proceedings of the Royal Society of London. Series B: Biological Sciences.