Environmental filtering of dense‐wooded species controls above‐ground biomass stored in African moist forests

Summary 1. Regional above-ground biomass estimates for tropical moist forests remain highly inaccurate mostly because they are based on extrapolations from a few plots scattered across a limited range of soils and other environmental conditions. When such conditions impact biomass, the estimation is biased. The effect of soil types on biomass has especially yielded controversial results. 2. We investigated the relationship between above-ground biomass and soil type in undisturbed moist forests in the Central African Republic. We tested the effects of soil texture, as a surrogate for soil resources availability and physical constraints (soil depth and hydromorphy) on biomass. Forest inventory data were collected for trees ‡20 cm stem diameter in 2754 0.5 ha plots scattered over 4888 km 2 . The plots contained 224 taxons, of which 209 were identified to species. Soil types were characterized from a 1:1 000 000 scale soil map. Species-specific values for wood density were extracted from the CIRAD’s data base of wood technological properties. 3. We found that basal area and biomass differ in their responses to soil type, ranging from 17.8 m 2 ha )1 (217.5 t ha )1 )t o 22.3 m 2 ha )1 (273.3 t ha )1 ). While shallow and hydromorphic soils support forests with both low stem basal area and low biomass, forests on deep resource-poor soils are typically low in basal area but as high in biomass as forests on deep resource-rich soils. We demonstrated that the environmental filtering of slow growing dense-wooded species on resource-poor soils compensates for the low basal area, and we discuss whether this filtering effect is due to low fertility or to low water reserve. 4. Synthesis. We showed that soil physical conditions constrained the amount of biomass stored in tropical moist forests. Contrary to previous reports, our results suggest that biomass is similar on resource-poor and resource-rich soils. This finding highlights both the importance of taking into account soil characteristics and species wood density when trying to predict regional patterns of biomass. Our findings have implications for the evaluation of biomass stocks in tropical forests, in the context of the international negotiations on climate change.

[1]  B. L. Welch ON THE COMPARISON OF SEVERAL MEAN VALUES: AN ALTERNATIVE APPROACH , 1951 .

[2]  E. Suzuki,et al.  Tap-root depth of tropical seedlings in relation to species-specific edaphic preferences , 2005, Journal of Tropical Ecology.

[3]  R. Condit,et al.  Demographic and life-history correlates for Amazonian trees , 2005 .

[4]  N. Higuchi,et al.  Variation in aboveground tree live biomass in a central Amazonian Forest: Effects of soil and topography , 2006 .

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

[6]  N. Picard,et al.  Manuel de référence pour l'installation de dispositifs permanents en forêt de production dans le Bassin du Congo , 2008 .

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

[8]  The Ecology of Trees in the Tropical Rain Forest , 2001 .

[9]  Robert L. Sanford,et al.  Nutrient Cycling in Moist Tropical Forest , 1986 .

[10]  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 .

[11]  Sandra A. Brown,et al.  Monitoring and estimating tropical forest carbon stocks: making REDD a reality , 2007 .

[12]  S. Titus,et al.  Relationships between tree slenderness coefficients and tree or stand characteristics for major species in boreal mixedwood forests , 1998 .

[13]  D. A. King,et al.  Soil‐related performance variation and distributions of tree species in a Bornean rain forest , 2005 .

[14]  J. Deckers,et al.  World Reference Base for Soil Resources , 1998 .

[15]  Y. Boulvert Etude géomorphologique de la République Centrafricaine : carte à 1/ 1000000 en deux feuilles ouest et est , 1996 .

[16]  David B. Clark,et al.  Landscape-scale variation in forest structure and biomass in a tropical rain forest , 2000 .

[17]  P. Couteron,et al.  Drawing ecological insights from a management-oriented forest inventory in French Guiana , 2003 .

[18]  R. Houghton,et al.  Aboveground Forest Biomass and the Global Carbon Balance , 2005 .

[19]  J. Slik,et al.  Soil nutrients affect spatial patterns of aboveground biomass and emergent tree density in southwestern Borneo , 2008, Oecologia.

[20]  H. A. Mooney,et al.  Maximum rooting depth of vegetation types at the global scale , 1996, Oecologia.

[21]  J. V. Soares,et al.  Distribution of aboveground live biomass in the Amazon basin , 2007 .

[22]  L. Poorter,et al.  Wood mechanics, allometry, and life-history variation in a tropical rain forest tree community. , 2006, The New phytologist.

[23]  D. L. Lauenstein,et al.  Coordination between leaf and stem traits related to leaf carbon gain and hydraulics across 32 drought-tolerant angiosperms , 2008, Oecologia.

[24]  C. Braak Canonical Correspondence Analysis: A New Eigenvector Technique for Multivariate Direct Gradient Analysis , 1986 .

[25]  Stephanie A. Bohlman,et al.  Importance of soils, topography and geographic distance in structuring central Amazonian tree communities , 2008 .

[26]  N. H. Ravindranath,et al.  2006 IPCC Guidelines for National Greenhouse Gas Inventories , 2006 .

[27]  Deutsche Ausgabe World Reference Base for Soil Resources 2006 , 2007 .

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

[29]  I. Brown,et al.  Uncertainty in the biomass of Amazonian forests: An example from Rondônia, Brazil , 1995 .

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

[31]  O. Phillips,et al.  Continental-scale patterns of canopy tree composition and function across Amazonia , 2006, Nature.

[32]  Richard A. Houghton,et al.  The spatial distribution of forest biomass in the Brazilian Amazon: a comparison of estimates , 2001 .

[33]  Peter J. Gregory Roots, rhizosphere and soil: the route to a better understanding of soil science? , 2006 .

[34]  K. Thompson,et al.  Integrated screening validates primary axes of specialisation in plants , 1997 .

[35]  P. Couteron,et al.  An operational, additive framework for species diversity partitioning and beta‐diversity analysis , 2007 .

[36]  C. Pendry,et al.  Altitudinal zonation of rain forest on Bukit Belalong, Brunei: soils, forest structure and floristics , 1997, Journal of Tropical Ecology.

[37]  P. Couteron,et al.  Regional variation in tropical forest tree species composition in the Central African Republic: an assessment based on inventories by forest companies , 2008, Journal of Tropical Ecology.

[38]  J. Terborgh,et al.  The regional variation of aboveground live biomass in old‐growth Amazonian forests , 2006 .

[39]  S. Gourlet‐Fleury,et al.  Can functional classification of tropical trees predict population dynamics after disturbance? , 2008 .

[40]  J. Laclau,et al.  A positive growth response to NaCl applications in Eucalyptus plantations established on K-deficient soils , 2010 .

[41]  J. Chave,et al.  Towards a Worldwide Wood Economics Spectrum 2 . L E a D I N G D I M E N S I O N S I N W O O D F U N C T I O N , 2022 .

[42]  J. Chambers,et al.  Relationship between soils and Amazon forest biomass: a landscape-scale study , 1999 .

[43]  Charles H. Cannon,et al.  Environmental correlates of tree biomass, basal area, wood specific gravity and stem density gradients in Borneo's tropical forests , 2010 .

[44]  P. Fearnside BRAZIL'S AMAZON FOREST AND THE GLOBAL CARBON PROBLEM , 1985 .

[45]  The spatial distribution of forest biomass in the Brazilian Amazon: a comparison of estimates , 2001 .

[46]  J. Plotkin,et al.  HABITAT PATTERNS IN TROPICAL RAIN FORESTS: A COMPARISON OF 105 PLOTS IN NORTHWEST BORNEO , 2002 .

[47]  Sandra A. Brown,et al.  State and change in carbon pools in the forests of tropical Africa , 1998 .

[48]  A. Grainger Difficulties in tracking the long-term global trend in tropical forest area , 2008, Proceedings of the National Academy of Sciences.

[49]  S. Goetz,et al.  Reply to Comment on ‘A first map of tropical Africa’s above-ground biomass derived from satellite imagery’ , 2008, Environmental Research Letters.

[50]  F. White The vegetation of Africa : a descriptive memoir to accompany the Unesco/AETFAT/UNSO vegetation map of Africa , 1985 .

[51]  T. Givnish Adaptive significance of evergreen vs. deciduous leaves : solving the triple paradox , 2002 .

[52]  Phillips,et al.  Changes in the carbon balance of tropical forests: evidence from long-term plots , 1998, Science.

[53]  G. Paoli,et al.  Soil nutrients and beta diversity in the Bornean Dipterocarpaceae: evidence for niche partitioning by tropical rain forest trees , 2006 .

[54]  S. Davies,et al.  Interspecific demographic trade‐offs and soil‐related habitat associations of tree species along resource gradients , 2007 .

[55]  B. Nicholas The ecology of trees in the tropical rain forest , 2004 .

[56]  Richard Condit,et al.  Error propagation and scaling for tropical forest biomass estimates. , 2004, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[57]  Y. Boulvert Carte phytogéographique de la République Centrafricaine (feuille Ouest-feuille Est) à 1:1000000 , 1986 .

[58]  Rattan Lal,et al.  Land Use, Land-Use Change and Forestry , 2015 .

[59]  J. P. Grime,et al.  Evidence for the Existence of Three Primary Strategies in Plants and Its Relevance to Ecological and Evolutionary Theory , 1977, The American Naturalist.

[60]  J. V. Soares,et al.  Distribution of aboveground live biomass in the Amazon basin , 2007 .

[61]  Ariel E. Lugo,et al.  Biomass Estimation Methods for Tropical Forests with Applications to Forest Inventory Data , 1989, Forest Science.

[62]  F. Humbel Caractérisation, par des mesures physiques, hydriques et d'enracinement, de sols de Guyane française à dynamique de l'eau superficielle , 1978 .

[63]  Kalle Ruokolainen,et al.  Dispersal, Environment, and Floristic Variation of Western Amazonian Forests , 2003, Science.

[64]  R. Bailey,et al.  Derivation, fitting, and implication of a compatible stem taper-volume-weight system for intensively managed, fast growing loblolly pine , 2002 .

[65]  F. S. Chapin,et al.  The Mineral Nutrition of Wild Plants , 1980 .

[66]  Mark S. Johnson,et al.  Relationships between soil hydrology and forest structure and composition in the southern Brazilian Amazon , 2007 .

[67]  Sean C. Thomas,et al.  Increasing carbon storage in intact African tropical forests , 2009, Nature.