UNDERSTANDING COLOMBIAN AMAZONIAN WHITE SAND FORESTS Maria Cristina Peñuela Mora The research presented in this thesis was carried out within the framework of the Plant Ecology and Biodiversity Group, Institute of Environmental Biology, Utrecht University and Ecology of Terrestrial Ecosystems Group, National University of Colombia. Maria Cristina Peñuela Mora Understanding Colombian Amazonian White Sand Forests Copyright © 2014, Maria Cristina Peñuela Mora All rights reserved ISBN: 978-90-393-6196-2 Keywords: Western Amazon, Terra firme forests, Forest dynamics, Drought, Seedling performance, Amazon caatinga Printed by: Panamericana Formas e Impresos S.A. Cover design: Camila Olarte Cover pictures: Mauritiella armata, Hevea nitida, marked trees and seedlings in the White sand forests, photos by Maria Cristina Peñuela. The tiny Dycimbe uaiparuensis seedling and Zafire Biological Station by Guillermo Santos. UNDERSTANDING COLOMBIAN AMAZONIAN WHITE SAND FORESTS ECOLOGIE VAN WITZANDBOSSEN IN COLOMBIA (met een samenvatting in het Nederlands) ENTENDIENDO LOS BOSQUES DE ARENAS BLANCAS DE LA AMAZONIA COLOMBIANA (con resumen en Español) Proefschrift ter verkrijging van de graad van doctor aan de Universiteit Utrecht op gezag van de rector magnificus, Prof. Dr. G.J. van der Zwaan, ingevolge het besluit van het college voor promoties in het openbaar te verdedigen op woensdag 1 oktober 2014 des middags te 12.45 uur door Maria Cristina Peñuela Mora geboren op 6 Juli te Bogotá, Colombia Promotor: Prof. Dr. R.G.A. Boot Co-promotor: Dr. H. ter Steege This thesis was written with financial support from the Netherlands Organization for International Higher Education (NUFFIC) and the Department of Science, Technology and Innovation, Colciencias, Colombia. To my son Luciano and my mother Elvira …and to my friends, women whose hearts and minds battle between their children, family and their studies. CONTENTS Chapter 1 General introduction 9 Chapter 2 Species composition of white sand forests in western 17 amazon Chapter 3 Eight years of forest dynamics in a white sand forest in 39 the Colombian Amazon – with reference to white sands and terra firme clay forests in the Amazon Chapter 4 Effects of soil conditions on the performance of tree seedlings 63 from terra firme and white sand forests – results of an in situ experiment transplanting soils and seedlings Chapter 5 Drought resistance of rainforest tree seedlings from white 85 sand and terra firme forests Chapter 6 Discussion 107 References 117 Resumen en español 140 English summary 146 Nederlandse samenvatting 151 Acknowledgements 156 Curriculum vitae 159 Publications 161 introduccion 8 – chapter 1 – 9 introduction GenerAl introdUCtion The Amazon region covers an area of nearly six million square kilometres (Ramos da Silva et al. 2008) and harbours the richest assemblage of plant species on Earth (Gentry 1988, de Oliveira & Mori 1999, ter Steege et al. 2000a, Hoorn & Wesselingh 2010, ter Steege et al. 2013). The diversity of plant species is partly due to the fact that the different species are restricted to a subset of environmental conditions (Fine et al. 2006). In particular, the species are restricted by physical soil characteristics such as: the soil water holding capacity, drainage, soil structure and nutrients, factors that affect tree growth, mortality and the floristic composition of tropical forests worldwide (Ferry et al. 2010, Quesada et al. 2009b). Amazonian forests have long been grouped in three broad categories: flooded forests, non-flooded or terra firme forests and white sand forests. White sand forests grow in large extensions in the Eastern Amazon and in Western Amazon they are located in small patches surrounded by terra firme forests that grow in different soils. Terra firme forests are found in clay soils and white sand forests grow in white sands or Podzols, which are also known as Spodozols or Arenosols (Quesada et al. 2011). Both types of soils are highly acidic, but terra firme soils have a higher cation exchange capacity and a higher amount of available total phosphorus. In general, white sand forests have extremely nutrient-poor soils that are strongly leached, and that are usually drained by black waters (Coomes & Grubb 1996, 1998a, Luizão et al. 2007, Quesada et al. 2009b, Quesada et al. 2011). In Podzols, complexes of aluminium, iron and organic compounds form a hardpan that impedes root penetration into deeper soil horizons (Schoenholtz et al. 2000) and causes waterlogging in the rainy season (Bongers et al. 1985, Medina et al. 1990, Quesada et al. 2011). In the dry season, the upper soil dries out due to the low water retention capacity (Duivenvoorden & Lips 1995, Williams et al. 2002), which increases mortality and decreases growth rates in tropical plants (Turner 1990, Condit et al. 1995, Veenendaal et al. 1995). Short periods of water stress are sufficient to cause important changes in the vegetation, not only in the floristic composition but also in the morphology and physiology of the plants (Franco & Dezzeo 1994, Hutyra et al. 2005, Walker et al. 2009). 10
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