Biogeosciences www.biogeosciences.net 1726-4170 1726-4189 6 9 2009 10.5194/bg-6-1883-2009 http://www.biogeosciences.net/6/1883/2009/ http://www.biogeosciences.net/6/1883/2009/bg-6-1883-2009.html http://www.biogeosciences.net/6/1883/2009/bg-6-1883-2009.pdf 1883 1902 2009-09-08 Influence of landscape heterogeneity on spatial patterns of wood productivity, wood specific density and above ground biomass in Amazonia L. O. Anderson liana.anderson@ouce.ox.ac.uk Y. Malhi R. J. Ladle L. E. O. C. Aragão Y. Shimabukuro O. L. Phillips T. Baker A. C. L. Costa J. S. Espejo N. Higuchi W. F. Laurance G. López-González A. Monteagudo P. Núñez-Vargas J. Peacock C. A. Quesada S. Almeida School of Geography and the Environment, University of Oxford, Oxford, UK Remote Sensing Division, National Institute for Space Research, São José dos Campos, Brazil Earth and Biosphere Institute, School of Geography, University of Leeds, LS2 9JT, England, UK Universidade Federal de Pará, Belém, Brazil Universidad Nacional San Antonia Abad del Cusco, Cusco, Perú Institito National de Pesquisas da Amazônia, Manaus, Brazil Smithsonian Tropical Research Institute, Balboa, Panama Herbario Vargas, Universidad Nacional San Antonio Abad del Cusco, Cusco, Perú Herbario Vargas, Universidad Nacional San Antonio Abad del Cusco, Cusco, Perú Museu Paraense Emilio Goeldi, Belém, Brazil Long-term studies using the RAINFOR network of forest plots have generated significant insights into the spatial and temporal dynamics of forest carbon cycling in Amazonia. In this work, we map and explore the landscape context of several major RAINFOR plot clusters using Landsat ETM+ satellite data. In particular, we explore how representative the plots are of their landscape context, and test whether bias in plot location within landscapes may be influencing the regional mean values obtained for important forest biophysical parameters. Specifically, we evaluate whether the regional variations in wood productivity, wood specific density and above ground biomass derived from the RAINFOR network could be driven by systematic and unintentional biases in plot location. Remote sensing data covering 45 field plots were aggregated to generate landscape maps to identify the specific physiognomy of the plots. In the Landsat ETM+ data, it was possible to spectrally differentiate three types of <i>terra firme</i> forest, three types of forests over Paleovarzea geomorphologycal formation, two types of bamboo-dominated forest, palm forest, <i>Heliconia</i> monodominant vegetation, swamp forest, disturbed forests and land use areas. Overall, the plots were generally representative of the forest physiognomies in the landscape in which they are located. Furthermore, the analysis supports the observed regional trends in those important forest parameters. This study demonstrates the utility of landscape scale analysis of forest physiognomies for validating and supporting the finds of plot based studies. Moreover, the more precise geolocation of many key RAINFOR plot clusters achieved during this research provides important contextual information for studies employing the RAINFOR database. Almeida-Filho, R. and Shimabukuro, Y. E.: Digital processing of a Landsat-TM time series for mapping and monitoring degraded areas caused by independent gold miners, Roraima State, Brazilian Amazon, Rem. Sens. Environ., 79, 42–50, 2002. Andreae, M. O., Jones, C. D., and Cox, P. M.: Strong present-day aerosol cooling implies a hot future, Nature, 435, 1187–1190, 2005. Aragão, L. E. O. C., Shimabukuro, Y. E., Espírito-Santo, F. D. B., and Williams, M.: Landscape pattern and spatial variability of leaf area index in Eastern Amazonia, Forest. Ecol. Manag., 211, 240–256, 2005. Aragão, L. E. O. C., Malhi, Y., Metcalfe, D. B., Silva-Espejo, J. E., Jiménez, E., Navarrete, D., Almeida, S., Costa, A. C. L., Salinas, N., Phillips, O. L., Anderson, L. O., Baker, T. R., Goncalvez, P. H., Huamán-Ovalle, J., Mamani-Solórzano, M., Meir, P., Monteagudo, A., Peñuela, M. C., Prieto, A., Quesada, C. A., Rozas-Dávila, A., Rudas, A., Silva Junior, J. A., and Vásquez, R.: Above- and below-ground net primary productivity across ten Amazonian forests on contrasting soils, Biogeosciences Discuss., 6, 2441–2488, 2009. Asner, G. P. and Wessman, C. A.: Scaling PAR absorption from the leaf to landscape level in spatially heterogeneous ecosystems, Ecol. Model., 103, 81–97 1997. Baker, T. R., Swaine, M. D., and Burslem, D. F.: Variation in tropical forest growth rates: combined effects of functional group composition and resource availability Perspectives in Plant Ecology, Evolution and Systematics, 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., Silva, J. N., and Martínez, R. V.: Increasing biomass in Amazonian forest plots, Philos. T. Roy. Soc. B, 359, 353–365, 2004a. Baker, T. R., Phillips, O. L., Malhi, Y., Almeida, S., Arroyo, L., Di Fiore, A., Killeen, T., Laurance, S. G., Laurance, W. F., Lewis, S. L., Lloyd, J., Monteagudo, A., Neill, D. A., Patiño, S., Pitman, N. C. A., Silva, N., and Vásquez Martínez, R.: Variation in wood density determines spatial patterns in Amazonian forest biomass, Global Change Biol., 10, 545–562, 2004b. Baker, T. R., Phillips, O. L., Laurance, W. F., Pitman, N. C. A., Almeida, S., Arroyo, L., DiFiore, A., Erwin, T., Higuchi, N., Killeen, T. J., Laurance, S. G., Nascimento, H., Monteagudo, A., Neill, D. A., Silva, J. N. M., Malhi, Y., López Gonzalez, G., Peacock, J., Quesada, C. A., Lewis, S. L., and Lloyd, J.: Do species traits determine patterns of wood production in Amazonian forests?, Biogeosciences, 6, 297–307, 2009. Betts, R. A., Cox, P. M., Collins, M., Harris, P. P., Huntingford, C., and Jones, C. D.: The role of ecosystem-atmosphere interactions in simulated Amazonian precipitation decrease and forest dieback under global climate warming, Theor. Appl. Climatol., 78, 137–158, 2004. Bicheron, P., Defourny, P., Brockmann, C., Schouten, L.,Vancutsem, C., Huc, M., Bontemps, S., Leroy, M., Achard, F., Herold, M., Ranera, F., and Arino, O.: Globcover, ESA / ESA GlobCover Project, available on-line: http://ionia1.esrin.esa.int/index.asp, 2008. Bins, L. S. E. G. J. and Fonseca, L. M. G.: Um método de classificação não supervisionada por regiões, in: Simpósio Brasileiro de Computação Gráfica e Processamento de Imagens, Recife, Brazil, 1, 65–68, 1993. Câmara, G., Souza, R. C. M., Freitas, U. M., and Garrido, J.: SPRING: Integrating remote sensing and GIS by object-oriented data modelling, Computers & Graphics, 20, 395–403, 1996. Chambers, J. Q., Santos, J., Ribeiro, R. J., and Higuchi, N.: Tree damage, allometric relationships, and above-ground net primary production in central Amazon forest, Forest. Ecol. Manag., 152, 73–84, 2001. Chave, J., Condit, R., Aguilar, S., Hernandez, A., Lao, S., and Perez, R.: Error propagation and scaling for tropical forest biomass estimates, Philos. T. Roy. Soc. B, 359, 409–420 2004. 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 forest, Oecologia, 145, 87–99, 2005. Chave, J., Navarrete, D., Almeida, S., \'Alvarez, E., Aragão, L. E. O. C., Bonal, D., Châtelet, P., Silva Espejo, J., Goret, J.-Y., von Hildebrand, P., Jiménez, E., Patiño, S., Peñuela, M. C., Phillips, O. L., Stevenson, P., and Malhi, Y.: Regional and temporal patterns of litterfall in tropical South America, Biogeosciences Discuss., 6, 7565–7597, 2009. 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. Clinebell II, R. R., Phillips, O. L., Gentry, A. H., Stark, N., and Zuuring, H.: Prediction of neotropical tree and liana species richness from soil and climatic data, Biodivers. Conserv., 4, 56–90, 1995. Condit, R., Pitman, N., Leigh Jr., E. G., Chave, J., Terborgh, J., Foster, R. B., Nunez, P., Aguilar, S., Valencia, R., Villa, G., Muller-Landau, H. C., Losos, E., and Hubbell, S. P.: Beta-diversity in tropical forest trees, Science, 295, 666–669, doi:10.1126/science.1066854, 2002. de Grandi, G. F., Mayaux, P., Malingreau, J. P., Rosenqvist, A., Saatchi, S., and Simard, M.: New perspectives on global ecosystems from wide-area radar mosaics: Flooded forest mapping in the tropics, Int. J. Remote Sens., 21, 1235–1249, 2000. Duque, A., Cavelier, J., and Posada, A.: Strategies of tree occupation at a local scale in terra firme forests in the Colombian Amazon, Biotropica, 35, 20–27, 2003. Fitzgerald, R. W. and Lees, B. G.: Assessing the classification accuracy of multisource remote sensing data, Rem. Sens. Environ., 47, 362–368, 1994. Foody, G. M. and Hill, R. A.: Classification of tropical forest classes from Landsat TM data, Int. J. Rem. Sens., 17, 2353–2367, 1996. Franklin, J. and Woodcock, C. E.: Multiscale vegetation data for the mountains of Southern California: spatial and categorical resolution, Scale in Remote Sensing and GIS, edited by: Goodchild, D. A. Q. a. M. F., CRC/Lewis 141–168 pp., 1997. Gentry, A. H.: Tree species richness of upper Amazonian forests, P. Natl. Acad. Sci.-Biol., 85, 156–159, 1988. Hess, L., Melack, J. M., Novo, E., Barbosa, C. C. F., and Gastil, M.: Dual-season mapping of wetland inundation and vegetation for the central Amazon basin, Rem. Sens. Environ., 87, 404–428, 2003. Higuchi, N., dos Santos, J., 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. Hill, R. A.: Image segmentation for humid tropical forest classification in Landsat TM data, Int. J. Remote Sens., 20, 1039–1044, 1999. Honorio Coronado, E. N., Baker, T. R., Phillips, O. L., Pitman, N. C. A., Pennington, R. T., Vásquez Martínez, R., Monteagudo, A., Mogollón, H., Dávila Cardozo, N., Ríos, M., García-Villacorta, R., Valderrama, E., Ahuite, M., Huamantupa, I., Neill, D. A., Laurance, W. F., Nascimento, H. E. M., Soares de Almeida, S., Killeen, T. J., Arroyo, L., Núñez, P., and Freitas Alvarado, L.: Integrating regional and continental scale comparisons of tree composition in Amazonian terra firme forests, Biogeosciences Discuss., 6, 1421–1451, 2009. Hoorn, C.: An environmental reconstruction of the palaeo-Amazon River system (Middle to late Miocene, NW Amazonia), Palaeogeography, Palaeoclimatology, Palaeoecology, 112, pp. 187–238, 1994 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. Huston, M.: Soil nutrients and tree species richness in Costa Rican forests, J. Biogeogr., 7, 147–157, 1980. Kai, L. and Muller, J. P.: Segmenting satellite imagery: A region growing scheme, Digest – International Geoscience and Remote Sensing Symposium (IGARSS), 1075–1078, 1991. Laurance, W. F., Laurance, S. G., Ferreira, L. V., Rankin-de Merona, J. M., Gascon, C., and Lovejoy, T. E.: Biomass collapse in Amazonian forest fragments, Science, 278, 1117–1118, 1997. Laurance, W. F., Ferreira, L. V., Rankin-de Merona, J. M., and Laurance, S. G.: Rain forest fragmentation and the dynamics of Amazonian tree communities, Ecology, 79, 2032–2040, 1998. Lewis, S. L., Phillips, O. L., Baker, T. R., Lloyd, J., Malhi, Y., Almeida, S., Higuchi, N., Laurance, W. F., Neill, D., Silva, N., Terborgh, J., Torres Lezama, A., Vásquez, M. R., Brown, S., Chave, J., Kuebler, C., N\'uñez V., P., and Vinceti, B.: Concerted changes in tropical forest structure and dynamics: evidence from 50 South American long-term plots, Philos. T. Roy. Soc. B, 359, 421–436, 2004. Malhi, Y., Phillips, O. L., Baker, T., Almeida, S., Fredericksen, T., Grace, J., Higuchi, N., Killeen, T., Laurance, W. F., Leano, C., Lloyd, J., Meir, P., Monteagudo, A., Neill, D., Nunez, P. V., Panfil, S. N., Pitman, N., Rudas, A., Salomao, R., Saleska, S., Silva, N., Silveira, M., Sombroek, W. G., Valencia, 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., Almeida, S., Alvarez, E., Arroyo, L., Chave, J., Czimczik, C., Di Fiore, A., Higuchi, N., Killeen, T., Laurance, S. G., Laurance, W. F., Lewis, S. L., Mercado, L. M., Monteagudo, A., Neill, D. A., Pitman, N. C. A., Quesada, C. A., Silva, J. N. M., Vásquez Martínez, R., Terborgh, J., Vinceti, B., and Lloyd, J.: The above-ground wood productivity and net primary productivity of 100 Neotropical forests, Global Change Biol., 10, 563–591, 2004. Malhi, Y., Wood, D., Baker, T. R., Wright, J., and Phillips, O.: regional variation of above-ground live biomass in old-growth Amazonian forests, Global Change Biol., 12, 1–32, 2006. Malhi, Y., Roberts, J. T., Betts, R. A., Killeen, T. J., Li, W., and Nobre, C. A.: Climate change, deforestation and the fate of the Amazon, Science, 319, 169–172, 2008. Marengo, J. A.: Characteristics and spatio-temporal variability of the Amazon River Basin Water Budget, Clim. Dynam., 24, 11–22, 2005. Muller-Landau, H. C.: Interspecific and inter-site variation in wood specific gravity of tropical trees, Biotropica, 36, 20–32, 2004. Mercado, L. M., Lloyd, J., Dolman, A. J., Sitch, S., and Patiño, S.: Modelling basin-wide variations in Amazon forest productivity – Part~1: Model calibration, evaluation and upscaling functions for canopy photosynthesis, Biogeosciences, 6, 1247–1272, 2009. Nebel, G., Kvist, L. P., Vanclay, J. K., and Vidaurre, H.: Forest dynamics in flood plain forests in the Peruvian Amazon: effects of disturbance and implications for management, Forest. Ecol. Manag., 150, 79–92, 2001. Nepstad, D. C., Verssimo, A., Alencar, A., Nobre, C., Lima, E., Lefebvre, P., Schlesinger, P., Potter, C., Moutinho, P., Mendoza, E., Cochrane, M., and Brooks, V.: Large-scale impoverishment of Amazonian forests by logging and fire, Nature, 398, 505–508, 1999. Patiño, S., Lloyd, J., Paiva, R., Baker, T. R., Quesada, C. A., Mercado, L. M., Schmerler, J., Schwarz, M., Santos, A. J. B., Aguilar, A., Czimczik, C. I., Gallo, J., Horna, V., Hoyos, E. J., Jimenez, E. M., Palomino, W., Peacock, J., Peña-Cruz, A., Sarmiento, C., Sota, A., Turriago, J. D., Villanueva, B., Vitzthum, P., Alvarez, E., Arroyo, L., Baraloto, C., Bonal, D., Chave, J., Costa, A. C. L., Herrera, R., Higuchi, N., Killeen, T., Leal, E., Luizão, F., Meir, P., Monteagudo, A., Neil, D., Núñez-Vargas, P., Peñuela, M. C., Pitman, N., Priante Filho, N., Prieto, A., Panfil, S. N., Rudas, A., Salomão, R., Silva, N., Silveira, M., Soares deAlmeida, S., Torres-Lezama, A., Vásquez-Martínez, R., Vieira, I., Malhi, Y., and Phillips, O. L.: Branch xylem density variations across the Amazon Basin, Biogeosciences, 6, 545–568, 2009. Peacock, J., Baker, T. R., Lewis, S. L., Lopez-Gonzalez, G., and Phillips, O. L.: The RAINFOR plot database: monitoring forest biomass and dynamics, J. Veg. Sci., 18, 542–553, 2007. Phillips, O. L., Hall, P., Gentry, A. H., Sawyer, A. S., and Vasquez, R.: Dynamics and species richness of tropical rain forests, P. Natl. Acad. Sci.-Biol., 91, 2805–2809, 1994. Phillips, O. L., Malhi, Y., Higuchi, N., Laurance, W. F., Nuñez Vargas, P., Vásquez Martinez, R., Laurance, S. G., Ferriera, L. V., Stern, M., Brown, S., and Grace, J.: Changes in the carbon balance of tropical forest: evidence from long-term plots, Science, 282, 439–442, 1998. Phillips, O. L., Malhi, Y., Vinceti, B., Baker, T., Lewis, S. L., Higuchi, N., Laurance, W. F., N\'uñez, V. P., Vásquez, M. R., Laurance, S. G., Ferreira, L. V., Stern, M., Brown, S., and Grace, J.: Changes in growth of tropical forests: evaluating potential biases, Ecol. Appl., 12, 576–587, 2002. Phillips, O. L., N\'uñez Vargas, P., Monteagudo, A. L., Peña Cruz, A., Chuspe Zans, M.-E., Sánchez, W.G., Yli-Halla, M., and Rose, S.: Habitat association among Amazonian tree species: a landscape-scale approach, J. Ecol., 91, 757–775, 2003. Phillips, O., Baker T. R., Arroyo, L., Higuchi, N., Killeen, T., Laurance, W. F., Lewis, S. L., Lloyd, J., Malhi, Y., Monteagudo, A., Neill, D., N\'uñez Vargas, P., Silva, N., 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., Palacios, W., Patiño, S., Pitman, N., Olivier, J., 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. Phillips, O. L., Aragão, L., Fisher, J. B., Lewis, S. L., Lloyd, J., López-González, G., Malhi, Y,. Monteagudo, A., Peacock, J., Quesada, C., van der Heijden, G., Almeida, S., Amaral, I., Arroyo, L., Aymard, G., Baker, T. R., Banki, O. S., Blanc, L., Bonal, D., Brando, P., Chave, J., de Oliveira, A. C., Dávila Cardozo, N., Espejo, J., Feldpausch, T., Aparecida Freitas, M., Higuchi, N., Jiménez, E., Meir, P., Mendoza B. C., Morel, A., Neill, D., Nepstad, D., Patiño, S., Peñuela, M. C., Prieto, A., Ramírez, F., Schwarz, M., Silveira, M., Sota, A., ter Steege, H., Stropp, J., Vásquez, R., Zelazowski, P., Alvarez Dávila, E., Andelman, S., Erwin, T., di Fiore, A., Chao, K.-J., Honorio, E., Keeling, H., Killeen, T., Laurance, W., Nascimento, H., Peña Cruz, A., Pitman, N., N\'uñez Vargas, P., Ramírez, H., Rudas, A., Salamão, R., Silva, N., Terborgh, J., and Torres, A.: Drought sensitivity of the Amazon rainforest, Science, 323: 1344–1347, 2009. Quesada, C. A., Lloyd, J., Anderson, L. O., Fyllas, N. M., Schwarz, M., and Czimczik, C. I.: Soils of amazonia with particular reference to the rainfor sites, Biogeosciences Discuss., 6, 3851–3921, 2009. Quesada, C. A., Lloyd, J., Schwarz, M., Baker, T. R., Phillips, O. L., Patiño, S., Czimczik, C., Hodnett, M. G., 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., Jimenez, E. M., Paiva, R., Vieira, I., Neill, D. A., Silva, N., 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., Killeen, T., Peña Cruz, A., Pitman, N., Núñez Vargas, 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., 6, 3993–4057, 2009a. Quesada, C. A., Lloyd, J., Anderson, L. O., Fyllas, N. M., Schwarz, M., and Czimczik, C. I.: Soils of amazonia with particular reference to the rainfor sites, Biogeosciences Discuss., 6, 3851–3921, 2009b. Richards, J. A.: Remote Sensing Digital Analysis: An lntroduction, New York: SpringerVerlag, pp. 133–154, 1993. Rodriguez-Yi, J. L., Shimabukuro, Y. E., and Rudorff, B. F. T.: Image segmentation for classification of vegetation using NOAA AVHRR data, Int. J. Remote Sens., 21, 167–172, 2000. Saatchi, S., Malhi, Y., Zutta, B., Buermann, W., Anderson, L. O., Araujo, A. M., Phillips, O. L., Peacock, J., ter Steege, H., Lopez Gonzalez, G., Baker, T., Arroyo, L., Almeida, S., Higuchi, N., Killeen, T., Monteagudo, A., Neill, D., Pitman, N., Prieto, A., Salomão, R., Silva, N., Vásquez Martínez, R., Laurance, W., and Ramírez, H. A.: Mapping landscape scale variations of forest structure, biomass, and productivity in Amazonia, Biogeosciences Discuss., 6, 5461–5505, 2009. Salo, J., Kalliola, R., Hakkinen, I., Makinen, Y., Niemela, P., Puhakka, M., and Colley, P. D.: River dynamics and the diversity of Amazon lowland forest, Nature, 322, 254–258, 1986. Salovaara, K. J., Cárdenas, G. G., and Tuomisto, H.: Forest classification in an Amazonian rainforest landscape using pteridophytes as indicator species, Ecography, 27, 689–700, 2004. Salovaara, K. J., Thessler, S., Malik, R. N., and Tuomisto, H.: Classification of Amazonian primary rain forest vegetation using Landsat ETM+ satellite imagery, Rem. Sens. Environ., 97, 39–51, 2005. Shlien, S. and Smith, A.: A rapid method to generate spectral theme classification of Landsat imagery, Rem. Sens. Environ., 4, 67–77, 1975. Stehman, S. V.: Estimating standard errors of accuracy assessment statistics under cluster sampling, Rem. Sens. Environ., 60, 258–269, 1997. 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. Telles, E. D. C., de Camargo, P. B., Martinelli, L. A., Trumbore, S. E., da Costa, E. S., Santos, J., Higuchi, N., and Oliveira, R. C.: Influence of soil texture on carbon dynamics and storage potential in tropical forest soils of Amazonia, Global Biogeochem. Cycles, 17, 1–9, 2003. ter Steege, H., Sabatier, S., Castellanos, H., van Andel, T., Duivenvoorden, J., de Oliveira, A. A., Ek, R. C., Lilwah, R., Maas, P. J. M., and Mori, S. A.: An analysis of Amazonian floristic composition, including those of the Guiana Shield, J. Tropical Ecol., 16, 801–828, 2000. ter Steege, H., Pitman, N. C. A., Phillips, O. L., Chave, J., Sabatier, D., Duque, A., Molino, J.-F., Prévost, M.-F., Spichiger, R., Castellanos, H., von Hildebrand, P., and Vásquez, R.: Continental-scale patterns of canopy tree composition and function across Amazonia, Nature, 443, 444–447, 2006. Tuomisto, H. and Ruokolainen, K.: Distribution of Pteridophyta and Melastomataceae along an edaphic gradient in an Amazonian rain forest, J. Veg. Sci., 5, 25–34, 1994. Tuomisto, H., Ruokolainen, K., and Yli-Halla, M.: Dispersal, environment, and floristic variation of western Amazonian forests, Science, 299, 241–244, doi:10.1126/science.1078037, 2003. Worbes, M.: The forest ecosystem of the floodplains, The Central Amazon Floodplain, edited by: Junk, W. J., Springer, Berlin, 223–266 pp., 1997. Wright, S. J.: Seasonal drought, soil fertility and the species density of tropical forest plant communities, Trends Ecol. Evol., 7, 260–263, 1992. Zonneveld, I.: The land unit: a fundamental concept on Landscape ecology, Landscape Ecol., 3, 67–89, 1989.