Biogeosciences www.biogeosciences.net 1726-4170 1726-4189 6 8 2009 10.5194/bg-6-1615-2009 http://www.biogeosciences.net/6/1615/2009/ http://www.biogeosciences.net/6/1615/2009/bg-6-1615-2009.html http://www.biogeosciences.net/6/1615/2009/bg-6-1615-2009.pdf 1615 1626 2009-08-11 After trees die: quantities and determinants of necromass across Amazonia K.-J. Chao O. L. Phillips T. R. Baker J. Peacock G. Lopez-Gonzalez R. Vásquez Martínez A. Monteagudo A. Torres-Lezama Earth and Biosphere Institute, School of Geography, University of Leeds, LS2 9JT, Leeds, UK International Master Program of Agriculture, College of Agriculture and Natural Resources, National Chung Hsing University, Taichung 40227, Taiwan Proyecto Flora del Perú, Jardín Botánico de Missouri, Oxapampa, Peru Herbario Vargas, Universidad Nacional San Antonio Abad del Cusco, Cusco, Peru INDEFOR, Facultad de Ciencias Forestales y Ambientales, Universidad de Los Andes, Mérida, Venezuela The Amazon basin, one of the most substantial biomass carbon pools on earth, is characterised by strong macroecological gradients in biomass, mortality rates, and wood density from west to east. These gradients could affect necromass stocks, but this has not yet been tested. This study aims to assess the stocks and determinants of necromass across Amazonian forests. Field-based and literature data were used to find relationships between necromass and possible determinants. Furthermore, a simple model was applied to estimate and extrapolate necromass stocks across terra firma Amazonian forests. In eight northwestern and three northeastern Amazonian permanent plots, volumes of coarse woody debris (≥10 cm diameter) were measured in the field and the density of each decay class was estimated. Forest structure and historical mortality data were used to determine the factors controlling necromass. Necromass is greater in forests with low stem mortality rates (northeast) rather than in forests with high stem mortality rates (northwest) (58.5±10.6 and 27.3±3.2 Mg ha−1, respectively). Using all published necromass values, we find that necromass across terra firma forests in Amazonia is positively related to both forest dynamics (mortality mass inputs and a surrogate for decomposition rate (average wood density of living trees)) and forest structure (biomass), but is better explained by forest dynamics. We propose an improved method to estimate necromass for plots where necromass has not been measured. The estimates, together with other actual measurements of necromass, were scaled-up to project a total Amazonian necromass of 9.6±1.0 Pg C. The ratio of necromass (on average weighted by forest region) to coarse aboveground biomass is 0.127. Overall, we find (1) a strong spatial trend in necromass in parallel with other macroecological gradients and (2) that necromass is a substantial component of the carbon pool in the Amazon. 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., Patiño, S., Pitman, N. C. A., Silva, J. N. M., and Vásquez Martínez, R.: Variation in wood density determines spatial patterns in Amazonian forest biomass, Global Change Biol., 10, 545–562, 2004. Baker, T. R., Honorio Coronado, E. N., Phillips, O. L., Martin, J., van der Heijden, G. M. F., Garcia, M., and Silva Espejo, J.: Low stocks of coarse woody debris in a southwest Amazonian forest, Oecologia, 152, 495–504, 2007. Brown, I. F., Martinelli, L. A., Thomas, W. W., Moreira, M. Z., Ferreira, C. A. C., and Victoria, R. A.: Uncertainty in the biomass of Amazonian forests: an example from Rondônia, Brazil, For. Ecol. Manag., 75, 175–189, 1995. Carey, E. V., Brown, S., Gillespie, A. J. R., and Lugo, A. E.: Tree mortality in mature lowland tropical moist and tropical lower montane moist forests of Venezuela, Biotropica, 26, 255–265, 1994. Chambers, J. Q., Higuchi, N., Schimel, J. P., Ferreira, L. V., and Melack, J. M.: Decomposition and carbon cycling of dead trees in tropical forests of the central Amazon, Oecologia, 122, 380–388, 2000. 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. Manag., 152, 73–84, 2001. Chao, K.-J., Phillips, O. L., and Baker, T. R.: Wood density and stocks of coarse woody debris in a northwestern Amazonian landscape, Can. J. For. Res., 38, 795–825, 2008a. Chao, K.-J., Phillips, O. L., Gloor, E., Monteagudo, A., Torres-Lezama, A., and Vásquez Martínez, R.: Growth and wood density predict tree mortality in Amazon forests, J. Ecol., 96, 281–292, 2008b. Chave, J., Andalo, C., Brown, S., Cairns, M. A., Chambers, J. Q., Eamus, D., Fölster, H., Fromard, F., Higuchi, N., Kira, T., Lescure, J.-P., Nelson, B. W., Ogawa, H., Puig, H., Riéra, 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.: Regional and phylogenetic variation of wood density across 2456 neotropical tree species, Ecol. Appl., 16, 2356–2367, 2006. Clark, D. A., Brown, S., Kicklighter, D. W., Chambers, J. Q., Thomlinson, J. R., Ni, J., and Holland, E. A.: Net primary production in tropical forests: an evaluation and synthesis of existing field data, Ecol. Appl., 11, 371–384, 2001. Cummings, D. L., Kauffman, J. B., Perry, D. A., and Hughes, R. F.: Aboveground biomass and structure of rainforests in the southwestern Brazilian Amazon, For. Ecol. Manag., 163, 293–307, 2002. Delaney, M., Brown, S., Lugo, A. E., Torres-Lezama, A., and Bello Quintero, N.: The distribution of organic carbon in major components of forests located in five life zones of Venezuela, J. Ecol., 13, 697–708, 1997. Delaney, M., Brown, S., Lugo, A. E., Torres-Lezama, A., and Quintero, N. B.: The quantity and turnover of dead wood in permanent forest plots in six life zones of Venezuela, Biotropica, 30, 2–11, 1998. Elias, M. and Potvin, C.: Assessing inter- and intra-specific variation in trunk carbon concentration for 32 neotropical tree species, Can. J. For. Res., 33, 1039–1045, 2003. FAO: Global Forest Resources Assessment (Food and Agriculture Organization of the United Nations), http://www.fao.org/forestry/site/fra2000report/en/, access: 27 September 2005, 2000. Gale, N.: The aftermath of tree death: coarse woody debris and the topography in four tropical rain forests, Can. J. For. Res., 30, 1489–1493, 2000. Gerwing, J. J.: Degradation of forests through logging and fire in the eastern Brazilian Amazon, For. Ecol. Manag., 157, 131–141, 2002. Harmon, M. E., Franklin, J. F., Swanson, F. J., Sollins, P., Gregory, S. V., Lattin, J. D., Anderson, N. H., Cline, S. P., Aumen, N. G., Sedell, J. R., Lienkaemper, G. W., Cromack, K. J., and Cummins, K. W.: Ecology of coarse woody debris in temperate ecosystems, Adv. Ecol. Res., 15, 133–302, 1986. Harmon, M. E. and Sexton, J.: Guidelines for Measurements of Woody Detritus in Forest Ecosystems, US Long Term Ecological Research Network Office Publication Seattle, Washington, 1996. Higuchi, N., Santos, J. D., Ribeiro, R. J., Minette, L., and Biot, Y.: Biomassa da parte aérea da vegetação da floresta tropical \'umida de terra-firme da Amazônia Brasileira, Acta Amazonica, 28, 153–166, 1998. Houghton, R. A., Lawrence, K. T., Hackler, J. L., and Brown, S.: The spatial distribution of forest biomass in the Brazilian Amazon: a comparison of estimates, Global Change Biol., 7, 731–746, 2001. Kauffman, J. B., Uhl, C., and Cummings, D. L.: Fire in the Venezuela Amazon 1: Fuel biomass and fire chemistry in the evergreen rainforest of Venezuela, Oikos, 53, 167–175, 1988. Keller, M., Palace, M., Asner, G. P., Pereira, R., and Silva, J. N. M.: Coarse woody debris in undisturbed and logged forests in the eastern Brazilian Amazon, Global Change Biol., 10, 784–795, 2004. Klinge, H.: Biomassa y materia orgánica del suelo in el ecosistema de la pluviselva centro-amazonica, Acta Cient. Venez., 24, 174–181, 1973. López-Gonzalez, G., Baker, T. R., Lewis, S. L., Peacock, J., and Phillips, O. L.: Functional Trait Database, RAINFOR (http://www.geog.leeds.ac.uk/projects/rainfor/), date of extraction: 8 February 2007, 2006. Lewis, S. L., Phillips, O. L., Sheil, D., Vinceti, B., Baker, T. R., Brown, S., Graham, A. W., Higuchi, N., Hilbert, D. W., Laurance, W. F., Lejoly, J., Malhi, Y., Monteagudo, A., N\'uñez Vargas, P., Sonk\.e, B., Supardi, M. N. N., Terborgh, J. W., and Vásquez Martínez, R.: Tropical forest tree mortality, recruitment and turnover rates: calculation, interpretation and comparison when census intervals vary, J. Ecol., 92, 929–944, 2004. Malhi, Y., Phillips, O. L., Lloyd, J., Baker, T. R., Wright, J., Almeida, S., Arroyo, L., Frederiksen, T., Grace, J., Higuchi, N., Killeen, T., Laurance, W. F., Leaño, C., Lewis, S. L., Meir, P., Monteagudo, A., Neill, D., N\'uñez Vargas, P., Panfil, S. N., Patiño, S., Pitman, N., Quesada, C. A., Rudas-Ll., A., Salomão, R., Saleska, S., Silva, N., Silveira, M., Sombroek, W. G., Valencia, R., Vásquez Martínez, R., Vieira, I. C. G., and Vinceti, B.: An international network to monitor the structure, composition and dynamics of Amazonian forests (RAINFOR), J. Veg. Sci., 13, 439–450, 2002. Malhi, Y., Wood, D., Baker, T. R., Wright, J., Phillips, O. L., Cochrane, T., Meir, P., Chave, J., Almeida, S., Arroyo, L., Higuchi, N., Killeen, T. J., Laurance, S. G., Laurance, W. F., Lewis, S. L., Monteagudo, A., Neill, D. A., N\'uñez Vargas, P., Pitman, N. C. A., Quesada, C. A., Salomão, R., Silva, J. N. M., Torres Lezama, A., Terborgh, J., Vásquez Martínez, R., and Vinceti, B.: The regional variation of aboveground live biomass in old-growth Amazonian forests, Global Change Biol., 12, 1107–1138, 2006. Martius, C.: Decomposition of wood, in: The Central Amazon Floodplain – Ecology of a Pulsing System, edited by: Junk, W. J., Springer, Germany, 267–276, 1997. Martius, C. and Bandeira, A. G.: Wood litter stocks in tropical moist forest in central Amazonia, Ecotropica, 4, 115–118, 1998. Nascimento, H. E. M. and Laurance, W. F.: Total aboveground biomass in central Amazonian rainforests: a landscape-scale study, For. Ecol. Manag., 168, 311–321, 2002. Olson, J. S.: Energy storage and the balance of producers and decomposers in ecological systems, Ecology, 44, 322–331, 1963. Palace, M., Keller, M., and Silva, H.: Necromass production: studies in undisturbed and logged Amazon forests, Ecol. Appl., 18, 873–884, 2008. Palace, M. W., Keller, M., Asner, G., Silva, J. N. M., and Passos, C.: Necromass in undisturbed and logged forests in the Brazilian Amazon, For. Ecol. Manag., 238, 309–318, 2007. Peacock, J., Baker, T. R., Lewis, S. L., López-Gonzalez, G., and Phillips, O. L.: The RAINFOR plot database: monitoring forest biomass and dynamics, J. Veg. Sci., 18, 535–542, 2007. Phillip, M. S.: Measuring Trees and Forests, 2nd ed., CAB International, Wallingford, UK, 1994. 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., N\'uñez Vargas, P., Silva, J. N. M., Terborgh, J., Vásquez Martínez, R., 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., Patiño, S., Pitman, N. C. A., Quesada, C. A., Saldias, M., Torres Lezama, A., and Vinceti, B.: Pattern and process in Amazon tree turnover, 1976–2001, Philos. T. Roy. Soc. B., 359, 381–407, 2004. RAINFOR: Amazon Forest Inventory Network, http://www.geog.leeds.ac.uk/projects/rainfor/, access: 8 February 2007. Rice, A. H., Hammond Pyle, E., Saleska, S. R., Hutyra, L., de Camargo, P. B., Portilho, K., Marques, D. F., and Wofsy, S. C.: Carbon balance and vegetation dynamics in an old-growth Amazonian forest, Ecol. Appl., 14(4) suppl., S55–S71, 2004. Saatchi, S. S., Houghton, R. A., Dos Santos Alvalá, R. C., Soares, J. V., and Yu, Y.: Distribution of aboveground live biomass in the Amazon basin, Global Change Biol., 13, 816–837, 2007. Saldarriaga, J. G., West, D. C., Tharp, M. L., and Uhl, C.: Long-term chronosequence of forest succession in the upper Rio Negro of Colombia and Venezuela, J. Ecol., 76, 938–958, 1988. Scott, D. A., Proctor, J., and Thompson, J.: Ecological studies on a lowland everygreen rain forest on Maracá Island, Roraima, Brasil. II. Litter and nutrient cycling, J. Ecol., 80, 705–717, 1992. Summers, P. M.: Estoque, decomposição, e nutrientes da liteira grossa em floresta de terra-firme, na Amazônia central, MS, Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil, 1998. Uhl, C., Buschbacher, R., and Serrão, E. A. S.: Abandoned pastures in eastern Amazônia, I: patterns and succession, J. Ecol., 76, 663–681, 1988. Uhl, C. and Kauffman, J. B.: Deforestation, fire susceptibility, and potential tree responses to fire in the eastern Amazon, Ecology, 71, 437–449, 1990. Vásquez Martínez, R.: Flórula de las Reservas Biológicas de Iquitos, Per\'u, Missouri Botanical Garden, St. Louis, 1997. Vásquez Martínez, R. and Phillips, O. L.: Allpahuayo: floristics, structure, and dynamics of a high-diversity forest in Amazonian Peru, Annals Missouri Botanical Garden, 87, 499–527, 2000. Veillon, J. P.: El crecimiento de algunos bosques naturales de Venezuela en relación con los parámetros del medio ambiente, Revista Forestal Venezolana, 29, 5–120, 1985.