Regional and phylogenetic variation of wood density across 2456 Neotropical tree species.

Wood density is a crucial variable in carbon accounting programs of both secondary and old-growth tropical forests. It also is the best single descriptor of wood: it correlates with numerous morphological, mechanical, physiological, and ecological properties. To explore the extent to which wood density could be estimated for rare or poorly censused taxa, and possible sources of variation in this trait, we analyzed regional, taxonomic, and phylogenetic variation in wood density among 2456 tree species from Central and South America. Wood density varied over more than one order of magnitude across species, with an overall mean of 0.645 g/cm3. Our geographical analysis showed significant decreases in wood density with increasing altitude and significant differences among low-altitude geographical regions: wet forests of Central America and western Amazonia have significantly lower mean wood density than dry forests of Central and South America, eastern and central Amazonian forests, and the Atlantic forests of Brazil; and eastern Amazonian forests have lower wood densities than the dry forests and the Atlantic forest. A nested analysis of variance showed that 74% of the species-level wood density variation was explained at the genus level, 34% at the Angiosperm Phylogeny Group (APG) family level, and 19% at the APG order level. This indicates that genus-level means give reliable approximations of values of species, except in a few hypervariable genera. We also studied which evolutionary shifts in wood density occurred in the phylogeny of seed plants using a composite phylogenetic tree. Major changes were observed at deep nodes (Eurosid 1), and also in more recent divergences (for instance in the Rhamnoids, Simaroubaceae, and Anacardiaceae). Our unprecedented wood density data set yields consistent guidelines for estimating wood densities when species-level information is lacking and should significantly reduce error in Central and South American carbon accounting programs.

[1]  G. B. Williamson,et al.  EXTREME RADIAL CHANGES IN WOOD SPECIFIC GRAVITY IN SOME TROPICAL PIONEERS , 2007 .

[2]  Ahmed Koubaa,et al.  Phenotypic Correlations Between Juvenile-Mature Wood Density and Growth in Black Spruce , 2007 .

[3]  S. Wright,et al.  Tropical forests in a changing environment. , 2005, Trends in ecology & evolution.

[4]  J. Chambers,et al.  Tree allometry and improved estimation of carbon stocks and balance in tropical forests , 2005, Oecologia.

[5]  A. Groover What genes make a tree a tree? , 2005, Trends in plant science.

[6]  Philip M. Fearnside,et al.  Wood density in dense forest in central Amazonia, Brazil , 2005 .

[7]  T. Speck,et al.  Plant growth forms: an ecological and evolutionary perspective. , 2005, The New phytologist.

[8]  Campbell O. Webb,et al.  Phylomatic: tree assembly for applied phylogenetics , 2005 .

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

[10]  A. Di Fiore,et al.  Variation in wood density determines spatial patterns inAmazonian forest biomass , 2004 .

[11]  J. Terborgh,et al.  The above‐ground coarse wood productivity of 104 Neotropical forest plots , 2004 .

[12]  Louis S. Santiago,et al.  Coordinated changes in photosynthesis, water relations and leaf nutritional traits of canopy trees along a precipitation gradient in lowland tropical forest , 2004, Oecologia.

[13]  H. Muller‐Landau Interspecific and Inter‐site Variation in Wood Specific Gravity of Tropical Trees , 2004 .

[14]  J. Chave,et al.  Structure and Biomass of Four Lowland Neotropical Forests , 2004 .

[15]  Daniel Sabatier,et al.  A spatial model of tree α-diversity and tree density for the Amazon , 2003, Biodiversity & Conservation.

[16]  J. Fisher,et al.  Why vines have narrow stems: Histological trends in Bauhinia (Fabaceae) , 1991, Oecologia.

[17]  J. Lundberg,et al.  An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants : APG II THE ANGIOSPERM PHYLOGENY GROUP * , 2003 .

[18]  S. Hubbell,et al.  GAP‐DEPENDENT RECRUITMENT, REALIZED VITAL RATES, AND SIZE DISTRIBUTIONS OF TROPICAL TREES , 2003 .

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

[20]  David R. Anderson,et al.  Model selection and multimodel inference : a practical information-theoretic approach , 2003 .

[21]  D. Woodcock,et al.  Does canopy position affect wood specific gravity in temperate forest trees? , 2003, Annals of botany.

[22]  Renske,et al.  A spatial model of tree α-diversity and -density for the Amazon , 2003 .

[23]  Campbell O. Webb,et al.  Phylogenies and Community Ecology , 2002 .

[24]  Eric R. Ziegel,et al.  Generalized Linear Models , 2002, Technometrics.

[25]  Stephen P. Hubbell,et al.  Beta-Diversity in Tropical Forest Trees , 2002, Science.

[26]  F. Meinzer Functional convergence in plant responses to the environment , 2002, Oecologia.

[27]  P. Parolin Radial gradients in wood specific gravity in trees of Central Amazonian floodplains , 2002 .

[28]  G. B. Williamson,et al.  Geographic Variation in Wood Specific Gravity: Effects of Latitude, Temperature, and Precipitation , 2002 .

[29]  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.

[30]  D. S. Hammond,et al.  CHARACTER CONVERGENCE, DIVERSITY, AND DISTURBANCE IN TROPICAL RAIN FOREST IN GUYANA , 2001 .

[31]  J. Terborgh,et al.  DOMINANCE AND DISTRIBUTION OF TREE SPECIES IN UPPER AMAZONIAN TERRA FIRME FORESTS , 2001 .

[32]  D. Woodcock Wood specific gravity of trees and forest types in the southern Peruvian Amazon , 2000 .

[33]  Martin Worbes,et al.  Wood density of trees in black water floodplains of Rio Jaú National Park, Amazonia, Brazil , 2000 .

[34]  M. Chase,et al.  A phylogenetic analysis of Rhamnaceae using rbcL and trnL-F plastid DNA sequences. , 2000, American journal of botany.

[35]  David R. Anderson,et al.  Model Selection and Inference: A Practical Information-Theoretic Approach , 2001 .

[36]  E. Suzuki Diversity in specific gravity and water content of wood among Bornean tropical rainforest trees , 1999, Ecological Research.

[37]  R. Burnham,et al.  The history of neotropical vegetation: New developments and status , 1999 .

[38]  L. Ferreira,et al.  Central Amazonian floodplains: effect of two water types on the wood density of trees. , 1998 .

[39]  Philip M. Fearnside,et al.  WOOD DENSITY FOR ESTIMATING FOREST BIOMASS IN BRAZILIAN AMAZONIA , 1997 .

[40]  A. Lugo,et al.  Estimating biomass and biomass change of tropical forests , 1997 .

[41]  Frederick N. Scatena,et al.  BACKGROUND AND CATASTROPHIC TREE MORTALITY IN TROPICAL MOIST, WET, AND RAIN FORESTS , 1996 .

[42]  R. Sederoff,et al.  Genetic mapping of quantitative trait loci controlling growth and wood quality traits in Eucalyptus grandis using a maternal half-sib family and RAPD markers. , 1996, Genetics.

[43]  J. Cornelius Heritabilities and additive genetic coefficients of variation in forest trees , 1994 .

[44]  V. Favrichon Classification des espèces arborées en groupes fonctionnels en vue de la réalisation d'un modèle de dynamique de peuplement en forêt Guyanaise , 1994 .

[45]  A. Lugo,et al.  Wood Densities of Tropical Tree Species , 1992 .

[46]  Brian D. Farrell,et al.  Escalation of Plant Defense: Do Latex and Resin Canals Spur Plant Diversification? , 1991, The American Naturalist.

[47]  G. B. Williamson,et al.  WOOD SPECIFIC GRAVITY GRADIENTS IN TROPICAL DRY AND MONTANE RAIN FOREST TREES , 1989 .

[48]  William M. Schaffer,et al.  Plant strategies and the dynamics and structure of plant communities , 1989 .

[49]  G. B. Williamson,et al.  Radial Gradients in the Specific Gravity of Wood in Some Tropical and Temperate Trees , 1989 .

[50]  A. Tyree,et al.  Vulnerability of Xylem to Cavitation and Embolism , 1989 .

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

[52]  Josefina Barajas-Morales Wood Specific Gravity in Species from Two Tropical Forests in Mexico , 1987 .

[53]  D. Hoekman,et al.  Ecological Wood Anatomy of the Woody Southern Californian Flora , 1985 .

[54]  C. Jordan,et al.  Succession and Nutrient Dynamics Following Forest Cutting and Burning in Amazonia , 1984 .

[55]  Martin Chudnoff,et al.  Tropical timbers of the world , 1984 .

[56]  G. B. Williamson Gradients in wood specific gravity of trees , 1984 .

[57]  A. Gentry,et al.  Neotropical Floristic Diversity: Phytogeographical Connections Between Central and South America, Pleistocene Climatic Fluctuations, or an Accident of the Andean Orogeny? , 1982 .

[58]  W. Hillis,et al.  Eucalypts for wood production. , 1978 .

[59]  S. Carlquist ECOLOGICAL FACTORS IN WOOD EVOLUTION: A FLORISTIC APPROACH , 1977 .

[60]  R. Archer,et al.  REACTION WOOD: INDUCTION AND MECHANICAL ACTION! , 1977 .

[61]  Peter H. Raven,et al.  Angiosperm Biogeography and Past Continental Movements , 1974 .

[62]  P. Raven,et al.  BUTTERFLIES AND PLANTS: A STUDY IN COEVOLUTION , 1964 .

[63]  K. Esau Anatomy of seed plants , 1960 .