species distribution; tropical trees; tropical Africa; environmental factors; Moran; statistics; spatial distribution; Cameroon; soil nutrients; central Africa
Résumé :
[en] Species distribution within plant communities results from both the influence of deterministic processes, related to environmental conditions, and neutral processes related to dispersal limitation and stochastic events, the relative importance of each factor depending on the observation scale.
Assessing the relative contribution of environment necessitates controlling for spatial dependences among data points. Recent methods, combining multiple regression and Moran's eigenvectors maps (MEM), have been proved successful in disentangling the influence of pure spatial processes related to dispersal limitation, pure environmental variables (not spatially structured) and spatially structured environmental properties. However, the latter influence is usually not testable when using advanced spatial models like MEM.
To overcome this issue, we propose an original approach, based on torus-translations and Moran spectral randomizations, to test the fraction of species abundance variation that is jointly explained by space and seven soil variables, using three environmental and tree species abundance data sets (consisting of 120, 52 and 34 plots of 0·2 ha each, located along 101-, 66- and 35-km-long transect-like inventories, respectively) collected in tropical moist forests in southern Cameroon.
The overall abundance of species represented by ≥30 individuals, and 27% of these species taken individually, were significantly explained by fine-scale (<5 km) and/or broad-scale (5–100 km) spatially structured variations in soil nutrient concentrations (essentially the concentration of available Mn, Mg and Ca) along the 120-plots area. The number of significant tests considerably decreased when investigating the two smaller data sets, which mostly resulted from low statistical power rather than weaker floristic and/or edaphic variation captured among plots.
Synthesis. Our results provide evidence that tree species turnovers are partly controlled by spatially structured concentrations in soil nutrients at scales ranging from few hundreds of metres to c. 100 km, a poorly documented subject in Central African forests. We also highlight the usefulness of our testing procedure to correctly interpret the space-soil fraction of variation partitioning analyses (which always accounted here for the most important part of the soil contribution), as this fraction was sometimes relatively high (R2 values up to c. 0·3) but nearly or not significant.
Disciplines :
Sciences de la terre & géographie physique Sciences de l’environnement & écologie Biologie végétale (sciences végétales, sylviculture, mycologie...)
Auteur, co-auteur :
Vleminckx, Jason
Doucet, Jean-Louis ; Université de Liège > Ingénierie des biosystèmes (Biose) > Laboratoire de Foresterie des régions trop. et subtropicales
Angiosperm Phylogeny Group (2009) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society, 161, 105–121.
Baldeck, C.A., Harms, K.E., Yavitt, J.B. et al. (2013) Soil resources and topography shape local tree community structure in tropical forests. Proceedings of the Royal Society, 280. doi:10.1098/rspb.2012.2532
Baldeck, C.A., Tupayachi, R., Sinca, F., Jaramillo, N. & Asner, G.P. (2016) Environmental drivers of tree community turnover in western Amazonian forests. Ecography, 39, 1089–1099.
Blanchet, F.G., Legendre, P. & Borcard, D. (2008) Forward selection of explanatory variables. Ecology, 89, 2623–2632.
Borcard, D., Gillet, F. & Legendre, P. (2011) Numerical Ecology with R. Springer, New York, NY, USA.
Borcard, D. & Legendre, P. (2002) All-scale spatial analysis of ecological data by means of principal coordinates of neighbour matrice. Ecological Modelling, 153, 51–68.
Chang, L.W., Zelený, D., Li, C.-F., Chiu, S.-T. & Hsieh, C.-F. (2013) Better environmental data may reverse conclusions about niche- and dispersal-based processes in community assembly. Ecology, 94, 2145–2151.
Chase, J.M. (2014) Spatial scale resolves the niche versus neutral theory debate. Journal of Vegetation Science, 25, 319–322.
Chuyong, G.B., Kenfack, D., Harms, K.E., Duncan, W.T., Condit, R. & Comita, L.S. (2011) Habitat specificity and diversity of tree species in an African wet tropical forest. Plant Ecology, 213, 1363–1374.
Condit, R., Engelbrecht, B.M.J., Pino, D., Perez, R. & Turner, B.L. (2013) Species distributions in response to individual soil nutrients and seasonal drought across a community of tropical trees. Proceedings of the National Academy of Sciences of the United States of America, 110, 5064–5068.
Dauby, G. & Hardy, O.J. (2011) Sampled-based estimation of diversity sensu stricto by transforming Hurlbert diversities into effective number of species. Ecography, 34, 001–012.
Deblauwe, V., Kennel, P. & Couteron, P. (2012) Testing pairwise association between spatially autocorrelated variables: a new approach using surrogate lattice data. PLoS ONE, 7, e48766.
Dray, S., Legendre, P. & Peres-Neto, P.R. (2006) Spatial modelling: a comprehensive framework for principal coordinate analysis of neighbour matrices (PCNM). Ecological Modelling, 196, 483–493.
Ezekiel, M. (1930) Methods of Correlation Analysis. Wiley & Sons, New York, NY, USA.
Fayolle, A., Engelbrecht, B., Freycon, V. et al. (2012) Geological substrates shape tree species and trait distributions in African moist forests. PLoS ONE, 7, e42381.
Fayolle, A., Loubota Panzou, G.J., Drouet, T., Swaine, M.D., Bauwens, S., Vleminckx, J., Biwolé, A., Lejeune, P. & Doucet, J.-L. (2016) Taller trees, denser stands and greater biomass in semi-deciduous than in evergreen lowland central African forests. Forest Ecology and Management, 374, 42–50.
Fine, P.V.A., Daly, D.C., Villa Muñoz, G., Mesones, I. & Cameron, K.M. (2005) The contribution of edaphic heterogeneity to the evolution and diversity of Burseraceae trees in the Western Amazon. Evolution, 59, 1464–1478.
Fox, J. & Weisberg, S. (2011) car: An {R} Companion to Applied Regression, 2nd edn. Sage, Thousand Oaks, CA, USA. Available at: http://socserv.socsci.mcmaster.ca/jfox/Books/Companion
Franqueville, A. (1973) Atlas régional Sud-Ouest I. Echelles 1/500000 et 1/1 000000. République Unie du Cameroun. ORSTOM, Paris, France.
Gause, I. (1934) The Struggle for Existence. Williams & Wilkins, Baltimore, MD, USA.
Gourlet-Fleury, S., Rossi, V., Réjou-Méchain, M. et al. (2011) Environmental filtering of dense-wooded species controls above-ground biomass stored in African moist forests. Journal of Ecology, 99, 981–990.
Griffith, D.A. & Peres-Neto, P.R. (2006) Spatial modeling in ecology: the flexibility of eigenfunction spatial analysis. Ecology, 87, 2603–2613.
Hardy, O.J. & Sonké, B. (2004) Spatial pattern analysis of tree species distribution in a tropical rain forest of Cameroon: assessing the role of limited dispersal and niche differentiation. Forest Ecology and Management, 197, 191–202.
Harms, K.E., Condit, R., Hubbell, S.P. & Foster, R.B. (2001) Habitat association of trees and shrubs in a 50-ha neotropical forest plot. Journal of Ecology, 89, 947–959.
Hubbell, S.P. (2001) The Unified Neutral Theory of Biodiversity and Biogeography. Princeton University Press, Princeton, NJ, USA.
Itoh, A., Ohkubo, T., Nanami, S., Tan, S. & Yamakura, T. (2010) Comparison of statistical tests for habitat associations in tropical forests: a case study of sympatric dipterocarp trees in a Bornean forest. Forest Ecology and Management, 259, 323–332.
Janzen, D.H. (1971) Herbivores and the number of tree species in tropical forests. The American Naturalist, 104, 501–528.
John, R., Dalling, J.W., Harms, K.E. et al. (2007) Soil nutrients influence spatial distributions of tropical tree species. Proceedings of the National Academy of Sciences of the United States of America, 104, 864–869.
Jones, A., Breuning-Madsen, H., Brossard, M. et al. (eds.) (2013) Soil Atlas of Africa. European Commission, Publications Office of the European Union, Luxembourg, Germany.
Lan, G., Hu, Y., Cao, M. & Zhu, H. (2011) Topography-related spatial distribution of dominant tree species in a tropical seasonal rain forest in China. Forest Ecology and Management, 262, 1507–1513.
Lebrun, J.-P. & Stork, A. (2008) Tropical African Flowering Plants. Ecology and Distribution, vol. 4 Fabaceae (Desmodium-Zornia). Conservatoires et Jardins botaniques de la ville de Genève, Genève, Switzerland.
Legendre, P. & Gallagher, E.D. (2001) Ecologically meaningful transformations for ordination of species data. Oecologia, 129, 271–280.
Legendre, P., Mi, X., Ren, H., Ma, K., Yu, M., Sun, I. & He, F. (2009) Partitioning beta diversity in a subtropical broad-leaved forest of China. Ecology, 90, 663–674.
Letouzey, R. (1985) Notice de la carte phytogéographique du Cameroun au 1/500.000. Institut de la Carte Internationale de la Végétation, Toulouse and Institut de la Recherche Agronomique, Yaoundé, Cameroon.
Millaleo, R., Reyes-Diaz, M., Ivanov, A.G., Mora, M.L. & Alberdi, M. (2010) Manganese as essential and toxic element for plants: transport, accumulation and resistance mechanisms. Journal of Soil Science and Plant Nutrition, 10, 470–481.
Naimi, B. (2013) usdm: Uncertainty analysis for species distribution models. R package version 1.1-12. http://CRAN.R-project.org/package=usdm
Newbery, D.M. & Proctor, J. (1984) Ecological studies in four contrasting lowland rain forests in Gunung Mulu National Park, Sarawak. IV. Association between tree distribution and soil factors. Journal of Ecology, 72, 475–493.
Newbery, D.M., Gartlan, J.S., McKey, D.B. & Waterman, P.G. (1986) The influence of drainage and soil phosphorus on the vegetation of Douala-Edea Forest Reserve, Cameroun. Vegetatio, 65, 149–162.
Ngachie, V. (1992) A general assessment of soil resources and soil fertility constraints in Cameroon on the basis of FAO-UNESCO soil map analysis. Tropicultura, 10, 61–63.
Oksanen, J., Blanchet, F.G., Kindt, R. et al. (2015) vegan: Community Ecology Package. R package version 2.2-1. http://CRAN.R-project.org/package=vegan
Pansu, M. & Gautheyrou, J. (2006) Handbook of Soil Analysis: Mineralogical, Organic and Inorganic Methods. Springer, Berlin, Germany.
Paoli, G.D. & Curran, L.M. (2007) Soil nutrients limit fine litter production and tree growth in mature lowland forest of southwestern Borneo. Ecosystems, 10, 503–518.
Paoli, G.D., Curran, L.M. & Zak, D.R. (2006) Soil nutrients and beta diversity in the Bornean Dipterocarpaceae: evidence for niche partitioning by tropical rain forest trees. Journal of Ecology, 94, 157–170.
Peres-Neto, P.R. & Legendre, P. (2010) Estimating and controlling for spatial structure in the study of ecological communities. Global Ecology and Biogeography, 2, 174–184.
Punchi-Manage, R., Wiegand, T., Wiegand, K., Getzin, S., Gunatilleke, C.V.S. & Gunatilleke, I.A.U.N. (2014) Effect of spatial processes and topography on structuring species assemblages in a Sri Lankan dipterocarp forest. Ecology, 95, 376–386.
Quesada, C.A., Phillips, O.L., Schwarz, M. et al. (2012) Basin-wide variations in Amazon forest structure and function are mediated by both soils and climate. Biogeosciences, 9, 2203–2246.
R Development Core Team (2014) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. Available at: http://www.R-project.org.
Reich, P.B., Ellsworth, D.S. & Uhl, C. (1995) Leaf carbon and nutrient assimilation and conservation in species of different successional status in an oligotrophic Amazonian forest. Functional Ecology, 9, 65–76.
Réjou-Méchain, M., Pélissier, R., Gourlet-Fleury, S., Couteron, P., Nasi, R. & Thompson, J.D. (2008) Regional variation in tropical forest tree species composition in the Central African Republic: an assessment based on inventories by forest companies. Journal of Tropical Ecology, 24, 663–674.
Robson, A.D. (ed.) (1989) Soil Acidity and Plant Growth. Academic Press, San Diego, CA, USA.
Russo, S.E., Davies, S.J., King, D.A. & Tan, S. (2005) Soil-related performance variation and distributions of tree species in a Bornean Rain Forest. Journal of Ecology, 93, 879–889.
Sabatier, D., Grimaldi, M., Prévost, M.-F., Guillaume, J., Godron, M., Dosso, M. & Curmi, P. (1997) The influence of soil cover organization on the floristic and structural heterogeneity of a Guianan Rain forest. Plant Ecology, 131, 81–108.
Sanchez, P.A. (1976) Properties and Management of Soils in the Tropics. Wiley, New York, NY, USA.
Sanchez, P.A., Couto, W. & Boul, S.W. (1982) The fertility capability classification system: Interpretation, applicability and modification. Geoderma, 27, 283–309.
Sollins, P. (1998) Factors influencing species composition in tropical lowland rain forest: does soil matter? Ecology, 79, 23–30.
Sterne, J.A. & Davey Smith, G. (2001) Sifting the evidence-what's wrong with significance tests? British Medical Journal, 322, 226–231.
Tuomisto, H., Ruokolainen, K. & Yli-Halla, M. (2003) Dispersal, environment, and floristic variation of Western Amazonian forests. Science, 299, 241–244.
Vennetier, P. (1963) Les hommes et leurs activités dans le nord du Congo-Brazzaville. PhD dissertation, University of Bordeaux, Bordeaux, France.
Vitousek, P.M. (1982) Nutrient cycling and nutrient use efficiency. The American Naturalist, 119, 553–572.
Vitousek, P.M. (2004) Nutrient Cycling and Limitation: Hawai'i as a Model System. Princeton University Press, Princeton, NJ, USA.
Vleminckx, J., Morin-Rivat, J., Biwolé, A.B., Daïnou, K., Gillet, J.-F., Doucet, J.-L., Drouet, T. & Hardy, O. (2014) Soil charcoal to assess the impacts of past human disturbances on tropical forests. PLoS ONE, 9, e108121.
Vleminckx, J., Drouet, T., Amani, C., Lisingo, J., Lejoly, J. & Hardy, O.J. (2015) Impact of fine-scale edaphic heterogeneity on tree species assembly in a central African rainforest. Journal of Vegetation Science, 26, 134–144.
Vleminckx, J., Doucet, J.-L., Morin-Rivat, J. et al. (2016) Data from: The influence of spatially structured soil properties on tree community assemblages at a landscape scale in the tropical forests of southern Cameroon. Dryad Digital Repository, http://dx.doi.org/10.5061/dryad.sp5dp.
Wagner, H.H. & Dray, S. (2015) Generating spatially constrained null models for irregularly spaced data using Moran spectral randomization methods. Methods in Ecology and Evolution, 6, 1169–1178.
WRB (2014) World Reference Base (WRB) for Soil Resources, 3rd edn. FAO, Rome, Italy.
Wright, S.J., Yavitt, J.B., Wurzburger, N. et al. (2011) Potassium, phosphorus, or nitrogen limit root allocation, tree growth, or litter production in a lowland tropical forest. Ecology, 92, 1616–1625.
