Convergent effects of elevation on functional leaf traits within and among species

Summary Spatial variation in filters imposed by the abiotic environment causes variation in functional traits within and among plant species. This is abundantly clear for plant species along elevational gradients, where parallel abiotic selection pressures give rise to predictable variation in leaf phenotypes among ecosystems. Understanding the factors responsible for such patterns may provide insight into the current and future drivers of biodiversity, local community structure and ecosystem function. In order to explore patterns in trait variation along elevational gradients, we conducted a meta-analysis of published observational studies that measured three key leaf functional traits that are associated with axes of variation in both resource competition and stress tolerance: leaf mass:area ratio (LMA), leaf nitrogen content per unit mass (Nmass) and N content per unit area (Narea). To examine whether there may be evidence for a genetic basis underlying the trait variation, we conducted a review of published results from common garden experiments that measured the same leaf traits. Within studies, LMA and Narea tended to decrease with mean annual temperature (MAT) along elevational gradients, while Nmass did not vary systematically with MAT. Correlations among pairs of traits varied significantly with MAT: LMA was most strongly correlated with Nmass and Narea at high-elevation sites with relatively lower MAT. The strengths of the relationships were equal or greater within species relative to the relationships among species, suggesting parallel evolutionary dynamics along elevational gradients among disparate biomes. Evidence from common garden studies further suggests that there is an underlying genetic basis to the functional trait variation that we documented along elevational gradients. Taken together, these results indicate that environmental filtering both selects locally adapted genotypes within plant species and constrains species to elevational ranges based on their ranges of potential leaf trait values. If individual phenotypes are filtered from populations in the same way that species are filtered from regional species pools, changing climate may affect both the species and functional trait composition of plant communities.

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

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

[3]  Melanie Smith,et al.  Alpine Treelines: Functional Ecology of the Global High Elevation Tree Limits , 2013 .

[4]  R. Shaw,et al.  ASSESSMENT OF PUBLICATION BIAS , 2013 .

[5]  P. Reich,et al.  New handbook for standardised measurement of plant functional traits worldwide , 2013 .

[6]  A. Agrawal,et al.  A Field Experiment Demonstrating Plant Life-History Evolution and Its Eco-Evolutionary Feedback to Seed Predator Populations , 2013, The American Naturalist.

[7]  M. Silman,et al.  Intra- and interspecific tree growth across a long altitudinal gradient in the Peruvian Andes. , 2012, Ecology.

[8]  P. Leadley,et al.  Impacts of climate change on the future of biodiversity. , 2012, Ecology letters.

[9]  C. Körner Alpine Treelines , 2012, Springer Basel.

[10]  Robert K. Colwell,et al.  Assessing the threat to montane biodiversity from discordant shifts in temperature and precipitation in a changing climate. , 2011, Ecology letters.

[11]  Y. Vitasse,et al.  To what extent is altitudinal variation of functional traits driven by genetic adaptation in European oak and beech? , 2011, Tree physiology.

[12]  C. Körner,et al.  Fine root traits in adult trees of evergreen and deciduous taxa from low and high elevation in the Alps , 2011, Alpine Botany.

[13]  Sebastian Schmidtlein,et al.  The role of plant functional trade-offs for biodiversity changes and biome shifts under scenarios of global climatic change , 2011 .

[14]  J. Abatzoglou,et al.  Changes in Climatic Water Balance Drive Downhill Shifts in Plant Species’ Optimum Elevations , 2011, Science.

[15]  Maja K. Sundqvist,et al.  Interactive effects of vegetation type and elevation on aboveground and belowground properties in a subarctic tundra , 2011 .

[16]  D. Wardle,et al.  Aboveground-Belowground Linkages: Biotic Interactions, Ecosystem Processes, and Global Change , 2010 .

[17]  Lucien Hoffmann,et al.  Intraspecific variability and trait‐based community assembly , 2010 .

[18]  F. Valladares,et al.  Global change and the evolution of phenotypic plasticity in plants , 2010, Annals of the New York Academy of Sciences.

[19]  B. Enquist,et al.  Opposing assembly mechanisms in a neotropical dry forest: implications for phylogenetic and functional community ecology. , 2009, Ecology.

[20]  L. Poorter,et al.  Causes and consequences of variation in leaf mass per area (LMA): a meta-analysis. , 2009, The New phytologist.

[21]  Jonathan M. Chase,et al.  Evolutionary diversification in stickleback affects ecosystem functioning , 2009, Nature.

[22]  L. Hedges,et al.  Introduction to Meta‐Analysis , 2009, International Coaching Psychology Review.

[23]  J. Craine Resource Strategies of Wild Plants , 2009 .

[24]  David D. Ackerly,et al.  Community assembly and shifts in plant trait distributions across an environmental gradient in coastal California , 2009 .

[25]  Nathan J B Kraft,et al.  Functional Traits and Niche-Based Tree Community Assembly in an Amazonian Forest , 2008, Science.

[26]  Campbell O. Webb,et al.  Are functional traits good predictors of demographic rates? Evidence from five neotropical forests. , 2008, Ecology.

[27]  P. Marquet,et al.  A Significant Upward Shift in Plant Species Optimum Elevation During the 20th Century , 2008, Science.

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

[29]  B. Beckage,et al.  A rapid upward shift of a forest ecotone during 40 years of warming in the Green Mountains of Vermont , 2008, Proceedings of the National Academy of Sciences.

[30]  A. Jarvis Hole-field seamless SRTM data, International Centre for Tropical Agriculture (CIAT) , 2008 .

[31]  Fernando Valladares,et al.  Ecological limits to plant phenotypic plasticity. , 2007, The New phytologist.

[32]  C. Körner The use of 'altitude' in ecological research. , 2007, Trends in ecology & evolution.

[33]  J. McElwain,et al.  Stomatal Frequency Change Over Altitudinal Gradients : Prospects for Paleoaltimetry , 2007 .

[34]  Michel Loreau,et al.  Eco‐evolutionary dynamics of communities and ecosystems , 2007 .

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

[36]  Brian J Enquist,et al.  Ecological and evolutionary determinants of a key plant functional trait: wood density and its community-wide variation across latitude and elevation. , 2007, American journal of botany.

[37]  J. Boughman Speciation in Sticklebacks , 2006 .

[38]  B. Potts,et al.  A framework for community and ecosystem genetics: from genes to ecosystems , 2006, Nature Reviews Genetics.

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

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

[41]  P. Reich,et al.  Fundamental trade-offs generating the worldwide leaf economics spectrum. , 2006, Ecology.

[42]  J. L. Parra,et al.  Very high resolution interpolated climate surfaces for global land areas , 2005 .

[43]  Tadashi Fukami,et al.  Long-term ecological dynamics: reciprocal insights from natural and anthropogenic gradients , 2005, Proceedings of the Royal Society B: Biological Sciences.

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

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

[46]  John Harte,et al.  INTEGRATING EXPERIMENTAL AND GRADIENT METHODS IN ECOLOGICAL CLIMATE CHANGE RESEARCH , 2004 .

[47]  C. Körner The nutritional status of plants from high altitudes , 1989, Oecologia.

[48]  C. Körner,et al.  Dry matter partitioning and root length/leaf area ratios in herbaceous perennial plants with diverse altitudinal distribution , 1987, Oecologia.

[49]  C. Körner,et al.  Altitudinal variation in stomatal conductance, nitrogen content and leaf anatomy in different plant life forms in New Zealand , 1986, Oecologia.

[50]  F. Woodward,et al.  Temperature-based population segregation in birch , 2003 .

[51]  G. Yohe,et al.  A globally coherent fingerprint of climate change impacts across natural systems , 2003, Nature.

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

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

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

[55]  Anurag A. Agrawal,et al.  Phenotypic Plasticity in the Interactions and Evolution of Species , 2001, Science.

[56]  A. Field Meta-analysis of correlation coefficients: a Monte Carlo comparison of fixed- and random-effects methods. , 2001, Psychological methods.

[57]  M. Wand Local Regression and Likelihood , 2001 .

[58]  Ülo Niinemets,et al.  GLOBAL-SCALE CLIMATIC CONTROLS OF LEAF DRY MASS PER AREA, DENSITY, AND THICKNESS IN TREES AND SHRUBS , 2001 .

[59]  S. Sultan Phenotypic plasticity for plant development, function and life history. , 2000, Trends in plant science.

[60]  Guohua Pan,et al.  Local Regression and Likelihood , 1999, Technometrics.

[61]  P. Reich,et al.  Convergence and correlations among leaf size and function in seed plants: a comparative test using independent contrasts. , 1999, American journal of botany.

[62]  C. Loehle Height growth rate tradeoffs determine northern and southern range limits for trees , 1998 .

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

[64]  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.

[65]  M. Pigliucci,et al.  Control of Phenotypic Plasticity Via Regulatory Genes , 1993, The American Naturalist.

[66]  D. Schluter,et al.  Ecological Character Displacement and Speciation in Sticklebacks , 1992, The American Naturalist.

[67]  W. Schlesinger,et al.  The global carbon dioxide flux in soil respiration and its relationship to vegetation and climate , 1992 .

[68]  Paul A. Keddy,et al.  Assembly and response rules: two goals for predictive community ecology , 1992 .

[69]  Christian Körner,et al.  Functional Morphology of Mountain Plants) , 1989 .

[70]  C. Körner,et al.  In situ photosynthetic responses to light, temperature and carbon dioxide in herbaceous plants from low and high altitude , 1987 .

[71]  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.

[72]  R. Huey Ecological Character Displacement in a Lizard , 1974 .

[73]  J. Clausen,et al.  Effect of varied environments on western North American plants , 1940 .