Plant functional type classifications in tropical dry forests in Costa Rica: leaf habit versus taxonomic approaches

Summary 1. One way to simplify the high taxonomic diversity of plant species in vegetation models is to place species into groups based on shared, dominant traits. Many studies have suggested that morphological and physiological traits of tropical dry forest tree species vary with leaf habit (i.e. leaves from evergreen, deciduous or semi-deciduous species) and thus this characteristic may serve as a useful way to distinguish ecologically meaningful functional types. 2. In this study we examine whether 10 plant traits vary with leaf habit in replicated leaves and individual trees of 87 species from a tropical dry forest in Costa Rica. We also looked for evidence of phylogenetic conservatism, i.e. closely related species sharing similar trait values compared to more distantly related taxa. 3. While some of the traits varied within and among individual trees of the same species, interspecific variation accounted for 57–83% of the variance among samples. Four traits in addition to leaf habit showed evidence of phylogenetic conservatism, but these results were strongly dependent on the inclusion of the 18 species of legumes (Fabaceae) in our dataset. Contrary to our predictions, none of the traits we measured differed among leaf habits. However, five traits (wood density, leaf C, leaf N, N ⁄ P and C ⁄ N) varied significantly between legumes and other functional types. Furthermore, when all high-nitrogen non-legume taxa were compared to the high-nitrogen legumes, six traits excluding leaf N differed significantly, indicating that legumes are functionally different from other tree species beyond high N concentrations. Similarly, the 18 legume taxa (which all have compound leaves) also differed from other compound-leaved species for six traits, thus leaf type does not explain these patterns. 4. Our main conclusions are that (i) a plant functional type classification based on leaf habit alone has little utility in the tropical dry forest we studied, and (ii) legumes have a different suite of traits including high leaf carbon and wood density in addition to high leaf nitrogen. Whether this result generalizes to other tropical forests is unknown, but merits future research due to the consequences of these traits for carbon storage and ecosystem processes.

[1]  H. Mooney,et al.  Photosynthetic Systems of Mediterranean-Climate Shrubs and Trees of California and Chile , 1970, The American Naturalist.

[2]  C. Delwiche,et al.  Nitrogen Isotope Distribution as a Presumptive Indicator of Nitrogen Fixation , 1979, Botanical Gazette.

[3]  G. Farquhar,et al.  Isotopic Composition of Plant Carbon Correlates With Water-Use Efficiency of Wheat Genotypes , 1984 .

[4]  P. G. Murphy,et al.  Ecology of Tropical Dry Forest , 1986 .

[5]  A. Gentry,et al.  Changes in Plant Community Diversity and Floristic Composition on Environmental and Geographical Gradients , 1988 .

[6]  A. Quispel Hellriegel and Wilfarth's discovery of (symbiotic) nitrogen fixation hundred years ago , 1988 .

[7]  M. Sobrado Cost-benefit relationships in deciduous and evergreen leaves of tropical dry forest species , 1991 .

[8]  R. Aerts The advantages of being evergreen. , 1995, Trends in ecology & evolution.

[9]  K. Killingbeck Nutrients in Senesced Leaves: Keys to the Search for Potential Resorption and Resorption Proficiency , 1996 .

[10]  J. A. Barone,et al.  HERBIVORY AND PLANT DEFENSES IN TROPICAL FORESTS , 1996 .

[11]  D. F. Grigal,et al.  NITROGEN MINERALIZATION AND PRODUCTIVITY IN 50 HARDWOOD AND CONIFER STANDS ON DIVERSE SOILS , 1997 .

[12]  Thomas M. Smith,et al.  Plant functional types : their relevance to ecosystem properties and global change , 1998 .

[13]  I. Brown,et al.  Stable carbon isotope ratio of tree leaves, boles and fine litter in a tropical forest in Rondônia, Brazil , 1998, Oecologia.

[14]  M. Béreau,et al.  Functional diversity in an Amazonian rainforest of French Guyana: a dual isotope approach (δ15N and δ13C) , 1998, Oecologia.

[15]  Eamus,et al.  Ecophysiological traits of deciduous and evergreen woody species in the seasonally dry tropics. , 1999, Trends in ecology & evolution.

[16]  A. Domenach,et al.  Leaf natural 15N abundance and total N concentration as potential indicators of plant N nutrition in legumes and pioneer species in a rain forest of French Guiana , 1999, Oecologia.

[17]  M. Kent,et al.  Plant functional types: an alternative to taxonomic plant community description in biogeography? , 2000 .

[18]  D. Bonal,et al.  Interspecific variability of δ13C among trees in rainforests of French Guiana: functional groups and canopy integration , 2000, Oecologia.

[19]  A. S. Evans,et al.  The Evolution of Plant Ecophysiological Traits: Recent Advances and Future Directions , 2000 .

[20]  Derek Eamus,et al.  Ecophysiology of trees of seasonally dry tropics: Comparisons among phenologies , 2001 .

[21]  S. Tarantola,et al.  Detecting vegetation leaf water content using reflectance in the optical domain , 2001 .

[22]  J. Singh,et al.  Effect of leaf habit and soil type on nutrient resorption and conservation in woody species of a dry tropical environment , 2001 .

[23]  Mark W. Chase,et al.  Evolution of the angiosperms: calibrating the family tree , 2001, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[24]  B. Enquist,et al.  Carbon isotope composition of tree leaves from Guanacaste, Costa Rica: comparison across tropical forests and tree life history , 2002, Journal of Tropical Ecology.

[25]  D. Bowman,et al.  Leaf attributes in the seasonally dry tropics: a comparison of four habitats in northern Australia , 2003 .

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

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

[28]  David D. Ackerly,et al.  FUNCTIONAL STRATEGIES OF CHAPARRAL SHRUBS IN RELATION TO SEASONAL WATER DEFICIT AND DISTURBANCE , 2004 .

[29]  P. Reich,et al.  Photosynthesis-nitrogen relations in Amazonian tree species , 1994, Oecologia.

[30]  Michael D. Abràmoff,et al.  Image processing with ImageJ , 2004 .

[31]  T. Gillespie,et al.  Diversity, composition, and structure of tropical dry forests in Central America , 2000, Plant Ecology.

[32]  N. Holbrook,et al.  Leaf physiology does not predict leaf habit; examples from tropical dry forest , 2005, Trees.

[33]  P. Reich,et al.  Global patterns of plant leaf N and P in relation to temperature and latitude. , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[34]  Jeannine Cavender-Bares,et al.  MULTIPLE TRAIT ASSOCIATIONS IN RELATION TO HABITAT DIFFERENTIATION AMONG 17 FLORIDIAN OAK SPECIES , 2004 .

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

[36]  William F. Fagan,et al.  Phylogenetic and Growth Form Variation in the Scaling of Nitrogen and Phosphorus in the Seed Plants , 2006, The American Naturalist.

[37]  R. DeFries,et al.  A global overview of the conservation status of tropical dry forests , 2006 .

[38]  Eric Garnier,et al.  Ecosystem productivity can be predicted from potential relative growth rate and species abundance. , 2006, Ecology letters.

[39]  Campbell O. Webb,et al.  Regional and phylogenetic variation of wood density across 2456 Neotropical tree species. , 2006, Ecological applications : a publication of the Ecological Society of America.

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

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

[42]  L. Puangchit,et al.  Contrasting seasonal leaf habits of canopy trees between tropical dry-deciduous and evergreen forests in Thailand. , 2006, Tree physiology.

[43]  Campbell O. Webb,et al.  Relationships among ecologically important dimensions of plant trait variation in seven neotropical forests. , 2007, Annals of botany.

[44]  H. G. Baker,et al.  SOIL AND STEM WATER STORAGE DETERMINE PHENOLOGY AND DISTRIBUTION OF TROPICAL DRY FOREST TREES ' , 2007 .

[45]  S. Lavorel,et al.  Incorporating plant functional diversity effects in ecosystem service assessments , 2007, Proceedings of the National Academy of Sciences.

[46]  Gregory P Asner,et al.  Controls over foliar N:P ratios in tropical rain forests. , 2007, Ecology.

[47]  Peter B Reich,et al.  Predicting leaf physiology from simple plant and climate attributes: a global GLOPNET analysis. , 2007, Ecological applications : a publication of the Ecological Society of America.

[48]  F. Bongers,et al.  Light-dependent leaf trait variation in 43 tropical dry forest tree species. , 2007, American journal of botany.

[49]  S. Cordell,et al.  Functional diversity of carbon-gain, water-use, and leaf-allocation traits in trees of a threatened lowland dry forest in Hawaii. , 2007, American journal of botany.

[50]  Andrew Hamilton,et al.  Stoichiometry and the New Biology: The Future Is Now , 2007, PLoS biology.

[51]  F. Valladares,et al.  Do we underestimate the importance of leaf size in plant economics? Disproportional scaling of support costs within the spectrum of leaf physiognomy. , 2007, Annals of botany.

[52]  L. Santiago,et al.  Extending the leaf economics spectrum to decomposition: evidence from a tropical forest. , 2007, Ecology.

[53]  Katherine N. Suding,et al.  Restoration through reassembly: plant traits and invasion resistance. , 2008, Trends in ecology & evolution.

[54]  M. Estiarte,et al.  Influence of water and terpenes on flammability in some dominant Mediterranean species , 2008 .

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

[56]  S. Pacala,et al.  Predictive Models of Forest Dynamics , 2008, Science.

[57]  Campbell O. Webb,et al.  Bioinformatics Applications Note Phylocom: Software for the Analysis of Phylogenetic Community Structure and Trait Evolution , 2022 .

[58]  J. Roy,et al.  High variation in foliage and leaf litter chemistry among 45 tree species of a neotropical rainforest community. , 2008, The New phytologist.

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

[60]  R. Pennington,et al.  Woody Plant Diversity, Evolution, and Ecology in the Tropics: Perspectives from Seasonally Dry Tropical Forests , 2009 .

[61]  J. Powers,et al.  Diversity and structure of regenerating tropical dry forests in Costa Rica: Geographic patterns and environmental drivers , 2009 .

[62]  Pete Smith,et al.  Integrating plant–soil interactions into global carbon cycle models , 2009 .

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

[64]  Nathan G. Swenson,et al.  Variation in leaf functional trait values within and across individuals and species: an example from a Costa Rican dry forest , 2010 .