The return of the variance: intraspecific variability in community ecology.

Despite being recognized as a promoter of diversity and a condition for local coexistence decades ago, the importance of intraspecific variance has been neglected over time in community ecology. Recently, there has been a new emphasis on intraspecific variability. Indeed, recent developments in trait-based community ecology have underlined the need to integrate variation at both the intraspecific as well as interspecific level. We introduce new T-statistics ('T' for trait), based on the comparison of intraspecific and interspecific variances of functional traits across organizational levels, to operationally incorporate intraspecific variability into community ecology theory. We show that a focus on the distribution of traits at local and regional scales combined with original analytical tools can provide unique insights into the primary forces structuring communities.

[1]  Márcio S Araújo,et al.  The ecological causes of individual specialisation. , 2011, Ecology letters.

[2]  S. Lavorel,et al.  On the importance of intraspecific variability for the quantification of functional diversity , 2012 .

[3]  N. Gotelli,et al.  NULL MODELS IN ECOLOGY , 1996 .

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

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

[6]  R. Callaway Positive interactions and interdependence in plant communities , 2007 .

[7]  F. Stuart Chapin,et al.  Functional Diversity — at the Crossroads between Ecosystem Functioning and Environmental Filters , 2007 .

[8]  Jon Norberg,et al.  A general multi-trait-based framework for studying the effects of biodiversity on ecosystem functioning. , 2007, Journal of theoretical biology.

[9]  Peter A. Abrams,et al.  The theory of limiting similarity , 1983 .

[10]  Mark Vellend,et al.  Ecological consequences of genetic diversity. , 2008, Ecology letters.

[11]  Louie H. Yang,et al.  The Ecology of Individuals: Incidence and Implications of Individual Specialization , 2002, The American Naturalist.

[12]  Cyrille Violle,et al.  Towards a trait-based quantification of species niche , 2009 .

[13]  L. Aarssen Ecological Combining Ability and Competitive Combining Ability in Plants: Toward a General Evolutionary Theory of Coexistence in Systems of Competition , 1983, The American Naturalist.

[14]  Mark Vellend,et al.  Conceptual Synthesis in Community Ecology , 2010, The Quarterly Review of Biology.

[15]  J. Kingsolver,et al.  Patterns and Power of Phenotypic Selection in Nature , 2007 .

[16]  J. P. Grime,et al.  The role of genotypic diversity in determining grassland community structure under constant environmental conditions , 2007 .

[17]  R M May,et al.  On the theory of niche overlap. , 1974, Theoretical population biology.

[18]  Wilfried Thuiller,et al.  Quantifying the relevance of intraspecific trait variability for functional diversity , 2011 .

[19]  Massimo Pigliucci,et al.  Phenotypic Plasticity: Beyond Nature and Nurture , 2001 .

[20]  Elizabeth E Crone,et al.  Causes and consequences of variation in plant population growth rate: a synthesis of matrix population models in a phylogenetic context. , 2010, Ecology letters.

[21]  J. Etterson,et al.  Plasticity to canopy shade in a monocarpic herb: within- and between-generation effects. , 2009, The New phytologist.

[22]  Jonathan Roughgarden,et al.  Niche Width: Biogeographic Patterns Among Anolis Lizard Populations , 1974, The American Naturalist.

[23]  Helmut Hillebrand,et al.  Biodiversity in a complex world: consolidation and progress in functional biodiversity research. , 2009, Ecology letters.

[24]  J. Dushoff,et al.  Intraspecific Variation and Species Coexistence , 2007, The American Naturalist.

[25]  T. Miki,et al.  Intraspecific niche flexibility facilitates species coexistence in a competitive community with a fluctuating environment , 2008 .

[26]  Janneke HilleRisLambers,et al.  High‐dimensional coexistence based on individual variation: a synthesis of evidence , 2010 .

[27]  R. Macarthur,et al.  The Limiting Similarity, Convergence, and Divergence of Coexisting Species , 1967, The American Naturalist.

[28]  Wilfried Thuiller,et al.  Defining and measuring ecological specialization , 2010 .

[29]  S. Schreiber,et al.  Why intraspecific trait variation matters in community ecology. , 2011, Trends in ecology & evolution.

[30]  J. Chave,et al.  Functional traits of individual trees reveal ecological constraints on community assembly in tropical rain forests , 2011 .

[31]  A. Minelli Phenotypic Integration: Studying the Ecology and Evolution of Complex Phenotypes , 2005 .

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

[33]  O. Hardy,et al.  Characterizing the phylogenetic structure of communities by an additive partitioning of phylogenetic diversity , 2007 .

[34]  B. Shipley From Plant Traits to Vegetation Structure: Chance and Selection in the Assembly of Ecological Communities , 2009 .

[35]  S. Wright,et al.  Evolution in Mendelian Populations. , 1931, Genetics.

[36]  James S. Clark,et al.  Resolving the biodiversity paradox. , 2007, Ecology letters.

[37]  W. Link,et al.  Individual Covariation in Life‐History Traits: Seeing the Trees Despite the Forest , 2002, The American Naturalist.

[38]  Jérémie Lecoeur,et al.  Competition, traits and resource depletion in plant communities , 2009, Oecologia.

[39]  J. P. Grime,et al.  Fine-scale community and genetic structure are tightly linked in species-rich grasslands , 2011, Philosophical Transactions of the Royal Society B: Biological Sciences.

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

[41]  H. Berg Cold Spring Harbor Symposia on Quantitative Biology.: Vol. LII. Evolution of Catalytic Functions. Cold Spring Harbor Laboratory, Cold Spring Harbor, NY, 1987, ISBN 0-87969-054-2, xix + 955 pp., US $150.00. , 1989 .

[42]  Cyrille Violle,et al.  Phylogenetic limiting similarity and competitive exclusion. , 2011, Ecology letters.

[43]  Paul A. Keddy,et al.  Assembly rules, null models, and trait dispersion : new questions from old patterns , 1995 .

[44]  M. Lechowicz,et al.  Alternative Designs and the Evolution of Functional Diversity , 2005, The American Naturalist.

[45]  Christopher Baraloto,et al.  Functional trait variation and sampling strategies in species-rich plant communities , 2010 .

[46]  Yi Ding,et al.  Within- and among-species variation in specific leaf area drive community assembly in a tropical cloud forest , 2011, Oecologia.

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

[48]  J. P. Grime,et al.  Community and ecosystem effects of intraspecific genetic diversity in grassland microcosms of varying species diversity. , 2010, Ecology.

[49]  Jonathan Roughgarden,et al.  Evolution of Niche Width , 1972, The American Naturalist.

[50]  Peter Chesson,et al.  Functional tradeoffs determine species coexistence via the storage effect , 2009, Proceedings of the National Academy of Sciences.

[51]  G. F. Gause The struggle for existence , 1971 .

[52]  Amy E. Miller,et al.  Niche complementarity due to plasticity in resource use: plant partitioning of chemical N forms. , 2010, Ecology.

[53]  Janneke HilleRisLambers,et al.  The importance of niches for the maintenance of species diversity , 2009, Nature.

[54]  S. Schreiber,et al.  The community effects of phenotypic and genetic variation within a predator population. , 2011, Ecology.

[55]  M. Mayfield,et al.  Opposing effects of competitive exclusion on the phylogenetic structure of communities. , 2010, Ecology letters.

[56]  L. Aarssen Competitive ability and species coexistence: a 'plant's-eye' view , 1989 .

[57]  B. McGill Exploring Predictions of Abundance from Body Mass Using Hierarchical Comparative Approaches , 2008, The American Naturalist.

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

[59]  S. Wright THE INTERPRETATION OF POPULATION STRUCTURE BY F‐STATISTICS WITH SPECIAL REGARD TO SYSTEMS OF MATING , 1965 .

[60]  William E. Stutz,et al.  Ecological release from interspecific competition leads to decoupled changes in population and individual niche width , 2010, Proceedings of the Royal Society B: Biological Sciences.

[61]  B. Oborny,et al.  Importance of plasticity and decision-making strategies for plant resource acquisition in spatio-temporally variable environments. , 2007, The New phytologist.

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

[63]  P. Chesson Mechanisms of Maintenance of Species Diversity , 2000 .

[64]  Selection ratios on community aggregated traits estimate ecological filters imposed on species by sites. , 2010, Ecology.

[65]  T. Dayan,et al.  Ecological and community‐wide character displacement: the next generation , 2005 .

[66]  O. Petchey,et al.  Including intraspecific variability in functional diversity. , 2009, Ecology.

[67]  R. Macarthur,et al.  The Theory of Island Biogeography , 1969 .

[68]  S. Pledger,et al.  A unified analysis of niche overlap incorporating data of different types , 2011 .

[69]  Evan Weiher,et al.  Advances, challenges and a developing synthesis of ecological community assembly theory , 2011, Philosophical Transactions of the Royal Society B: Biological Sciences.

[70]  J. Soussana,et al.  Effects of land-use change on productivity depend on small-scale plant species diversity. , 2009 .

[71]  Michael D. Collins,et al.  Supporting Online Material Materials and Methods Figs. S1 to S3 Tables S1 and S2 References Plant Genotypic Diversity Predicts Community Structure and Governs an Ecosystem Process , 2022 .

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

[73]  C. Schlichting,et al.  Variation: A Central Concept in Biology , 2007 .

[74]  N. Swenson,et al.  Functional trait assembly through ecological and evolutionary time , 2009, Theoretical Ecology.

[75]  Christopher Baraloto,et al.  Fine‐scale Microhabitat Heterogeneity in a French Guianan Forest , 2010 .

[76]  M. Vellend The consequences of genetic diversity in competitive communities. , 2006, Ecology.

[77]  C. Stern CONCLUDING REMARKS OF THE CHAIRMAN , 1950 .

[78]  IR-04-071 Adaptive Speciation : Epilogue , 2004 .

[79]  S. Higgins,et al.  TRY – a global database of plant traits , 2011, Global Change Biology.

[80]  P. Reich,et al.  A handbook of protocols for standardised and easy measurement of plant functional traits worldwide , 2003 .

[81]  Nathan J B Kraft,et al.  Response to Comment on “Functional Traits and Niche-Based Tree Community Assembly in an Amazonian Forest” , 2009, Science.

[82]  A. Ostling,et al.  Comment on “Functional Traits and Niche-Based Tree Community Assembly in an Amazonian Forest” , 2009, Science.

[83]  F. Bello,et al.  Community trait response to environment: disentangling species turnover vs intraspecific trait variability effects , 2011 .

[84]  L. V. Valen,et al.  MORPHOLOGICAL VARIATION AND WIDTH OF ECOLOGICAL NICHE , 1965 .

[85]  Martin Daufresne,et al.  A user's guide to functional diversity indices , 2010 .

[86]  G. Vieilledent,et al.  Intra-specific variability and the competition–colonisation trade-off: coexistence, abundance and stability patterns , 2010, Theoretical Ecology.

[87]  Eric Garnier,et al.  Consistency of species ranking based on functional leaf traits. , 2001, The New phytologist.

[88]  K. Holsinger,et al.  Genetics in geographically structured populations: defining, estimating and interpreting FST , 2009, Nature Reviews Genetics.

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

[90]  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.

[91]  Amy E. Miller,et al.  Plant uptake of inorganic and organic nitrogen: neighbor identity matters. , 2007, Ecology.

[92]  M. C. Ferrari,et al.  Intraspecific trait variants determine the nature of interspecific interactions in a habitat-forming species. , 2011, Ecology.

[93]  J. Cavender-Bares,et al.  The merging of community ecology and phylogenetic biology. , 2009, Ecology letters.

[94]  Marc T. J. Johnson,et al.  Community genetics: what have we accomplished and where should we be going? , 2011, Philosophical Transactions of the Royal Society B: Biological Sciences.

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

[96]  James S. Clark,et al.  References and Notes Supporting Online Material Individuals and the Variation Needed for High Species Diversity in Forest Trees , 2022 .

[97]  J Norberg,et al.  Phenotypic diversity and ecosystem functioning in changing environments: A theoretical framework , 2001, Proceedings of the National Academy of Sciences of the United States of America.

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

[99]  J. Ellers,et al.  Trait plasticity in species interactions: a driving force of community dynamics , 2010, Evolutionary Ecology.

[100]  R M May,et al.  Niche overlap as a function of environmental variability. , 1972, Proceedings of the National Academy of Sciences of the United States of America.