Biogeosciences www.biogeosciences.net 1726-4170 1726-4189 6 2 2009 10.5194/bg-6-297-2009 http://www.biogeosciences.net/6/297/2009/ http://www.biogeosciences.net/6/297/2009/bg-6-297-2009.html http://www.biogeosciences.net/6/297/2009/bg-6-297-2009.pdf 297 307 2009-02-25 Do species traits determine patterns of wood production in Amazonian forests? T. R. Baker t.r.baker@leeds.ac.uk O. L. Phillips W. F. Laurance N. C. A. Pitman S. Almeida L. Arroyo A. DiFiore T. Erwin N. Higuchi T. J. Killeen S. G. Laurance H. Nascimento A. Monteagudo D. A. Neill J. N. M. Silva Y. Malhi G. López Gonzalez J. Peacock C. A. Quesada S. L. Lewis J. Lloyd Earth and Biosphere Institute, School of Geography, University of Leeds, Leeds, UK Smithsonian Tropical Research Institute, Balboa, Panama Center for Tropical Conservation, Duke University, Durham, USA Museu Paraense Emilio Goeldi, Belém, Brazil Museo Noel Kempff Mercado, Santa Cruz, Bolivia Department of Anthropology, New York University, NY, USA Smithsonian Institution, Washington DC, USA Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil Center for Applied Biodiversity Science, Conservation International, Washington DC, USA Biological Dynamics of Forest Fragments Project, Manaus, Brazil Proyecto Flora del Perú, Jardin Botanico de Missouri, Oxapampa, Perú Missouri Botanical Garden, c/o Naturaleza y Cultura Internacional, Loja, Ecuador Center for International Forestry Research, Tapajos, Brazil EMBRAPA Amazonia Oriental, Belém, Brazil Oxford University Centre for the Environment, Oxford, UK Understanding the relationships between plant traits and ecosystem properties at large spatial scales is important for predicting how compositional change will affect carbon cycling in tropical forests. In this study, we examine the relationships between species wood density, maximum height and above-ground, coarse wood production of trees ≥10 cm diameter (CWP) for 60 Amazonian forest plots. Average species maximum height and wood density are lower in Western than Eastern Amazonia and are negatively correlated with CWP. To test the hypothesis that variation in these traits causes the variation in CWP, we generate plot-level estimates of CWP by resampling the full distribution of tree biomass growth rates whilst maintaining the appropriate tree-diameter and functional-trait distributions for each plot. These estimates are then compared with the observed values. Overall, the estimates do not predict the observed, regional-scale pattern of CWP, suggesting that the variation in community-level trait values does not determine variation in coarse wood productivity in Amazonian forests. Instead, the regional gradient in CWP is caused by higher biomass growth rates across all tree types in Western Amazonia. Therefore, the regional gradient in CWP is driven primarily by environmental factors, rather than the particular functional composition of each stand. These results contrast with previous findings for forest biomass, where variation in wood density, associated with variation in species composition, is an important driver of regional-scale patterns in above-ground biomass. Therefore, in tropical forests, above-ground wood productivity may be less sensitive than biomass to compositional change that alters community-level averages of these plant traits. Baker, T. R., Swaine, M. D., and Burslem, D. F. R. P.: Variation in tropical forest growth rates: combined effects of functional group composition and resource availability, Perspectives in Plant Ecology, Evol. Syst., 6, 21–36, 2003. Baker, T. R., Phillips, O. L., Malhi, Y., Almeida, S., Arroyo, L., Di Fiore, A., Erwin, T., Higuchi, N., Killeen, T. J., Laurance, S. G., Laurance, W. F., Lewis, S. L., Monteagudo, A., Neill, D. A., Vargas, P. N., Pitman, N. C. A., Silva, J. N. M., and Martinez, R. V.: Increasing biomass in Amazonian forest plots, Philos. Trans. R. Soc. Lond. Ser. B-Biol. Sci., 359, 353–365, 2004a. Baker, T. R., Phillips, O. L., Malhi, Y., Almeida, S., Arroyo, L., Di Fiore, A., Erwin, T., Killeen, T. J., Laurance, S. G., Laurance, W. F., Lewis, S. L., Lloyd, J., Monteagudo, A., Neill, D. A., Patino, S., Pitman, N. C. A., Silva, J. N. M., and Martinez, R. V.: Variation in wood density determines spatial patterns in Amazonian forest biomass, Global Change Biol., 10, 545–562, 2004b. Balvanera, P., Kremen, C., and Martineez-Ramos, M.: Applying community structure analysis to ecosystem function: examples from pollination and carbon storage, Ecol. Appl., 15, 360–375, 2005. Barberis, I. M. and Tanner, E. V. J.: Gaps and root trenching increase tree seedling growth in Panamanian semi-evergreen forest, Ecology, 86, 667–674, 2005. Bunker, D. E., DeClerck, F., Bradford, J. C., Colwell, R. K., Perfecto, I., Phillips, O. L., Sankaran, M., and Naeem, S.: Species loss and aboveground carbon storage in a tropical forest, Science, 310, 1029–1031, 2005. Chambers, J. Q., dos Santos, J., Ribeiro, R. J., and Higuchi, N.: Tree damage, allometric relationships, and above-ground net primary production in central Amazon forest, For. Ecol. Manage., 5348, 1–12, 2000. Chambers, J. Q., Higuchi, N., Teixeira, L. M., dos Santos, J., Laurance, S. G., and Trumbore, S. E.: Response of tree biomass and wood litter to disturbance in a Central Amazon forest, Oecologia, 141, 596–611, 2004. Chave, J.: Study of structural, successional and spatial patterns in tropical rain forests using TROLL, a spatially explicit forest model, Ecol. Model., 124, 233–254, 1999. Chave, J., Andalo, C., Brown, S., Cairns, M. A., Chambers, J. Q., Eamus, D., Folster, H., Fromard, F., Higuchi, N., Kira, T., Lescure, J. P., Nelson, B. W., Ogawa, H., Puig, H., Riera, B., and Yamakura, T.: Tree allometry and improved estimation of carbon stocks and balance in tropical forests, Oecologia, 145, 87–99, 2005. Chave, J., Muller-Landau, H. C., Baker, T. R., Easdale, T. A., ter Steege, H., and Webb, C. O.: Phylogenetic conservatism and regional variation in wood density among neotropical tree species, Ecol. Appl., 16, 2356–2367, 2006. Dixon, R. K., Brown, S., Houghton, R. A., Solomon, A. M., Trexler, M. C., and Wisniewski, J.: Carbon pools and flux of global forest ecosystems, Science, 263, 185–190, 1994. Enquist, B., West, G. B., Charnov, E. L., and Brown, J. H.: Allometric scaling of production and life-history variation in vascular plants, Nature, 401, 907–911, 1999. Falster, D. S. and Westoby, M.: Alternative height strategies among 45 dicot rain forest species from tropical Queensland, Australia, J. Ecol., 93, 521–535, 2005. Galwey, N. W.: Introduction to Mixed Modelling: Beyond Regression and Analysis of Variance. Wiley, Chichester, England, Hoboken, NJ, USA, 376 pp, 2006. Garnier, E., Cortez, J., Billès, G., Navas, M.-L., Roumet, C., Debussche, M., Laurent, G., Blanchard, A., Aubry, D., Bellmann, A., Neill, C., and Toussaint, J.-P.: Plant functional markers capture ecosystem properties during secondary succession, Ecology, 85, 2630–2637, 2004. Gentry, A. H. and Emmons, L. H.: Geographical variation in fertility, phenology, and composition of the understory of Neotropical forests, Biotropica, 19, 216–227, 1987. Hooper, D. U., Chapin, F. S., Ewel, J. J., Hector, A., Inchausti, P., Lavorel, S., Lawton, J. H., Lodge, D. M., Loreau, M., Naeem, S., Schmid, B., Setala, H., Symstad, A. J., Vandermeer, J., and Wardle, D. A.: Effects of biodiversity on ecosystem functioning: A consensus of current knowledge, Ecol. Monogr., 75, 3–35, 2005. Keeling, H. C., Baker, T. R., Vasquez Martinez, R., Monteagudo, A., and Phillips, O. L.: Contrasting patterns of diameter and biomass increment across tree functional groups in Amazonian forests, Oecologia, 158, 521–534, 2008. Killeen, T. J., Garcia E., E., and Beck, S. G.: Guia de arboles de Bolivia, Missouri Botanic Garden, St. Louis, USA, 958 pp., 1993. LaFrankie, J. V., Ashton, P. S., Chuyong, G., Co, L., Condit, R., Davies, S. J., Foster, R. B., Hubbell, S. P., Kenfack, D., Lagunzad, D., Losos, E. C., Nor, N. S. M., Tan, S., Thomas, D. W., Valencia, R., and Villa, G.: Contrasting structure and composition of the understorey in species-rich tropical rainforests, Ecology, 87, 2298–2305, 2006. Laurance, W. F., Oliveira, A. A., Laurance, S. G., Condit, R., Nascimento, H. E. M., Sanchez-Thorin, A. C., Lovejoy, T. E., Andrade, A., D'Angelo, S., Ribeiro, J. E., and Dick, C. W.: Pervasive alteration of tree communities in undisturbed Amazonian forests, Nature, 428, 171–175, 2004. Lavorel, S. and Garnier, E.: Predicting changes in community composition and ecosystem functioning from plant traits: revisiting the Holy Grail, Func. Ecol., 16, 545–556, 2002. Levine, J. M.: Species diversity and biological invasions: relating local process to community pattern, Science, 288, 852-854, 2000. Lewis, S. L. and Tanner, E. V. J.: Effects of above- and belowground competition on growth and survival of rain forest tree seedlings, Ecology, 81, 2525–2538, 2000. Lieberman, D. and Lieberman, M.: Forest tree growth and dynamics at La Selva, Costa Rica (1969–1982), J. Trop. Ecol., 3, 347–358, 1987. Malhi, Y., Phillips, O. L., Baker, T., Almeida, S., Fredericksen, T., Grace, J., Higuchi, N., Killeen, T., Laurance, W. L., Leaño, C., Lloyd, J., Meir, P., Monteagudo, A., Neill, D., N\'uñez Vargas, P., Panfil, S., Pitman, N., Rudas Ll, A., Salamão, R., Saleska, S., Silva, N., Silveira, M., Sombroek, W. G., Valencia, R., Vásquez Martínez, R., Vieira, I., and Vinceti, B.: An international network to understand the biomass and dynamics of Amazonian forests (RAINFOR), J. Veg. Sci., 13, 439–450, 2002. Malhi, Y., Baker, T. R., Phillips, O. L., Almeida, S., Alvarez, E., Arroyo, L., Chave, J., Czimczik, C. I., Di Fiore, A., Higuchi, N., Killeen, T. J., Laurance, S. G., Laurance, W. F., Lewis, S. L., Montoya, L. M. M., Monteagudo, A., Neill, D. A., Vargas, P. N., Patino, S., Pitman, N. C. A., Quesada, C. A., Salomao, R., Silva, J. N. M., Lezama, A. T., Martinez, R. V., Terborgh, J., Vinceti, B., and Lloyd, J.: The above-ground coarse wood productivity of 104 Neotropical forest plots, Global Change Biol., 10, 563–591, 2004. Patiño, S., Lloyd, J., Paiva, R., Quesada, C. A., Baker, T. R., Santos, A. J. B., Mercado, L. M., Malhi, Y., Phillips, O.L., Aguilar, A., Alvarez, E., Arroyo, L., Bonal, D., Costa, A. C. L., Czimczik, C. I., Gallo, J., Herrera, R., Higuchi, N., Horna, V., Hoyos, E. J., Jiménez, E. M., Killeen, T., Leal, E., Luizão, F., Meir, P., Monteagudo, A., Neill, D. A., N\'uñez-Vargas, P., Palomino, W., Peacock, J., Peña-Cruz, A., Peñuela, M. C., Pitman, N. C. A., Priante Filho, N., Prieto, A., Panfil, S. N., Rudas, A., Salomão, R., Silva, N. M., Silveira, M., Soares de Almeida, S., Torres-Lezama, A., Turriago, J. D., Vásquez, R., Schwarz, M., Sota, A., Schmerler, J., Vieira, I., Villanueva, B. and Vitzthum, P.: Branch xylem density variations across the Amazon Basin, Biogeosciences Discuss., 5, 2003–2047, 2008. Peacock, J., Baker, T. R., Lewis, S. L., Lopez-Gonzalez, G., and Phillips, O. L.: The RAINFOR database: monitoring forest biomass and dynamics, J. Veg. Sci., 18, 535–542, 2007. Phillips, O. L., Vásquez, R., Arroyo, L., Baker, T. R., Killeen, T., Lewis, S., Malhi, Y., Monteagudo, A., Neill, D., N\'uñez Vargas, P., Alexiades, M., Cerón, C., Di Fiore, A., Erwin, T., Jardim, A., Palacios, W., Saldias, M., and Vinceti, B.: Increasing dominance of large lianas in Amazonian forests, 2002. Phillips, O. L., Martinez, R. V., Vargas, P. N., Monteagudo, A. L., Zans, M. E. C., Sanchez, W. G., Cruz, A. P., Timana, M., Yli-Halla, M., and Rose, S.: Efficient plot-based floristic assessment of tropical forests, J. Trop. Ecol., 19, 629–645, 2003. Phillips, O. L., Baker, T. R., Arroyo, L., Higuchi, N., Killeen, T. J., Laurance, W. F., Lewis, S. L., Lloyd, J., Malhi, Y., Monteagudo, A., Neill, D. A., Vargas, P. N., Silva, J. N. M., Terborgh, J., Martinez, R. V., Alexiades, M., Almeida, S., Brown, S., Chave, J., Comiskey, J. A., Czimczik, C. I., Di Fiore, A., Erwin, T., Kuebler, C., Laurance, S. G., Nascimento, H. E. M., Olivier, J., Palacios, W., Patino, S., Pitman, N. C. A., Quesada, C. A., Salidas, M., Lezama, A. T., and Vinceti, B.: Pattern and process in Amazon tree turnover, 1976–2001, Philos. Trans. R. Soc. Lond. Ser. B-Biol. Sci., 359, 381–407, 2004. Pitman, N. C. A., Terborgh, J. W., Silman, M. R., N\'uñez, P. V., Neill, D. A., Cerón, C., Palacios, W. A., and Aulestia, M.: Dominance and distribution of tree species in upper Amazonian terra firme forests, Ecology, 82, 2101–2117, 2001. Pitman, N. C. A., Cerón, C., Reyes, C. I., Thurber, M., and Arellano, J.: Catastrophic natural origin of a species-poor tree community in the world's richest forest, J. Trop. Ecol., 21, 559–568, 2006. Poorter, L., Hawthorne, W., Bongers, F., and Sheil, D.: Maximum size distributions in tropical forest communities: relationships with rainfall and disturbance, J. Ecol., 96, 495–504, 2008a. Poorter, L., Wright, S. J., Paz, H., Ackerly, D. D., Condit, R., Ibarra-Manriquez, G., Harms, K. E., Licona, J. C., Martinez-Ramos, M., Mazer, S. J., Muller-Landau, H., Pena-Claros, M., Webb, C. O., and Wright, I. J.: Are functional traits good predictors of demographic rates? Evidence from five neotropical forests, Ecology, 89, 1908–1920, 2008b. Quesada, C. A., Lloyd, J., Schwarz, M., Baker, T. R., Phillips, O. L., Patiño, S., Czimczik, C. I., Hodnett, M., Herrera, R., Arneth, A., Lloyd, G., Malhi, Y., Dezzeo, N., Luizão, F. J., Santos, A. J. B., Schmerler, J., Arroyo, L., Silveira, M., Priante Filho, N., Jiménez, E. M., Paiva, R., Vieira, I., Neill, D. A., Silva, N. M., Peñuela, M. C., Monteagudo, A., Vásquez, R., Prieto, A., Rudas, A., Almeida, S., Higuchi, N., Lezama, A. T., López-González, G., Peacock, J., Fyllas, N. M., Alvarez Dávila, E., Erwin, T., di Fiore, A., Chao, K.-J., Honorio, E. N., Killeen, T. J., Peña Cruz, A., Pitman, N. C. A., N\'uñez, P., Salomão, R., Terborgh, J. and Ramírez, H.: Regional and large-scale patterns in Amazon forest structure and function are mediated by variations in soil physical and chemical properties, Biogeosciences Discuss., accepted, 2009. Ribeiro, J. E. L. d. S., Hopkins, M. J. G., Vicentini, A., Sothers, C. A., Costa, M. A. d. S., Brito, J. M. d., Souza, M. A. D. d., Martins, L. H. P., Lohmann, L. G., Assunção, P. A. C. L., Pereira, E. d. C., Silva, C. F. d., Mesquita, M. R., and Procópio, L. C.: Flora da Reserva Ducke, INPA-DFID, 799 pp., 1999. Santiago, L. S., Goldstein, G., Meinzer, F. C., Fisher, J. B., Machado, K., Woodruff, D., and Jones, T.: Leaf photosynthetic traits scale with hydraulic conductivity and wood density in Panamanian forest canopy trees, Oecologia, 140, 543–550, 2004. Sombroek, W.: Amazon landforms and soils in relation to biological diversity, Acta Amazonica, 30, 81–100, 2000. Sombroek, W. G.: Spatial and temporal patterns of Amazon rainfall, Ambio, 30, 388–396, 2001. Suding, K. N., Lavorel, S., Chapin, F. S., Cornelissen, J. H. C., Diaz, S., Garnier, E., Goldberg, D., Hooper, D. U., Jackson, S. T., and Navas, M.-L.: Scaling environmental change through the community-level: a trait-based response-and-effect framework for plants, Global Change Biol., 14, 1125–1140, 2008. Swaine, M. D. and Becker, P.: Woody life-form composition and association on rainfall and soil fertility gradients in Ghana, Plant Ecol., 145, 167–173, 1999. ter Steege, H., Pitman, N., Phillips, O. L., Chave, J., Sabatier, D., Duque, A., Molino, J.-F., Prévost, M.-F., Spichiger, R., Castellanos, H., von Hildebrand, P., and Martinez, R. V.: Continental scale patterns of canopy tree composition and function across Amazonia, Nature, 443, 444–447, 2006. Thomas, S. C. and Bazzaz, F. A.: Asymptotic height as a predictor of photosynthetic characteristics in Malaysian forest trees, Ecology, 80, 1607–1622, 1999. Turner, I. M.: The Ecology of Trees in the Tropical Rain Forest, Cambridge University Press, Cambridge, 2001. Vásquez Martínez, R.: Flórula de las Reservas Biológicas de Iquitos, Per\'u, Monographs in Systematic Botany from the Missouri Botanical Garden, edited by: Lleras, A. R. and Taylor, C. M., Missouri Botanical Garden, St. Louis, USA, 1046 pp., 1997. Vilà, M., Vayreda, J., Gracia, C., and Ibanez, J. J.: Does tree diversity increase wood production in pine forests?, Oecologia, 135, 299–303, 2003. Vilà, M., Vayreda, J., Comas, L., Ibáñez, J., Mata, T., and Obón, B.: Species richness and wood production: a positive association in Mediteranean forests, Ecol. Lett., 10, 241–250, 2007. Wardle, D. A., Bonner, K. I., and Barker, G. M.: Stability of ecosystem properties in response to above-ground functional group richness and composition, Oikos, 89, 11–23, 2000.