T OWARDS AN UNDERSTANDING OF TREE DIVERSITY A IN MAZONIAN FORESTS The research presented in this thesis was carried out at the Ecology & Biodiversity Group of the Institute of Environmental Science, Utrecht University, with financial support from: Project for the Advancement of Networked science in Amazonia “PAN-AMAZONIA” - EU Sixth Framework Programme (2002-2006) Programa BECA – ‘Instituto Internacional de Educação do Brasil’ and Gordon and Betty Moore Foundation National Council of Scientific and Technological Development (CNPq), Ministry of Science and Technology, Brazil Copyrights © 2011, Juliana Stropp All rights reserved. Cover design: Tito Fernandes Picture cover: Canopy view of a terra-firme forest at Pico da Neblina National Park, São Gabriel da Cachoeira, Brazil (Photograph by Peter van der Sleen) Printing: Proefschriftmaken.nl ISBN: 978-90-393-5526-8 T OWARDS AN UNDERSTANDING OF TREE DIVERSITY IN AMAZONI AN FORESTS OORZAKEN VAN BOOMDIVERSITEIT IN DE AMAZONE (met een samenvatting in het Nederlands) ENTENDENDO DIVERSIDADE DE ÁRVORES DAS FLORESTAS DA AMAZÔNIA (com resumo em Português) Proefschrift ter verkrijging van de graad van doctor aan de Universiteit Utrecht op gezag van de rector magnificus, prof. dr. J.C. Stoof, ingevolge het besluit van het college voor promoties in het openbaar te verdedigen op maandag 21 maart 2011 des middags te 4.15 uur door Juliana Stropp Carneiro geboren op 5 december 1975 te Campinas, Brazilie Promotor: Prof.dr. M. J. A. Werger Co-promotor: Dr. Hans ter Steege Para minha mãe, Marilde Stropp C ONTENTS 1 INTRODUCTION .......................................................................................... 9 2 DISENTANGLING REGIONAL AND LOCAL TREE DIVERSITY IN AMAZONIA ..................... 17 3 TREE COMMUNITIES OF WHITE-SAND AND TERRA-FIRME FORESTS OF THE UPPER RIO NEGRO ..................................................................................... 31 4 SEEDLING TRAITS AND HERBIVORY IN WHITE-SAND AND TERRA-FIRME FORESTS: AN ANALYSIS ACROSS A RESOURCE GRADIENT IN THE UPPER RIO NEGRO, BRAZIL ........ 47 5 COMPARING TREE COMMUNITIES OF WHITE-SAND AND TERRA-FIRME FORESTS ACROSS THREE AMAZONIAN REGIONS ............................................................. 71 6 DISCUSSION AND PERSPECTIVES .................................................................... 95 REFERENCES ............................................................................................. 105 APPENDIX 1: TREE SPECIES REGISTERED IN THE UPPER RIO NEGRO .......................... 121 APPENDIX 2: ATTRIBUTES OF TREE INVENTORY PLOTS .......................................... 135 APPENDIX 3: REFERENCES TO PLOT DATA ......................................................... 143 SUMMARY ............................................................................................... 145 SAMENVATTING ......................................................................................... 149 RESUMO .................................................................................................. 153 ACKNOWLEDGMENTS .................................................................................. 157 CURRICULUM VITAE .................................................................................... 161 LIST OF PUBLICATIONS ................................................................................. 163 AFFILIATIONS OF CO-AUTHORS ....................................................................... 165 1 I NTRODUCTION Amazonian forests arguably harbor the highest biodiversity of all terrestrial ecosystems on Earth. Both the origin of this extraordinary biodiversity and its current distribution are recently becoming better understood (ter Steege et al., 2003, Hoorn and Wesselingh, 2010). Processes such as long-term landscape evolution and climate change have been identified as important drivers of speciation and extinction, thus shaping the current patterns of biodiversity in Amazonian forests (Hoorn et al., 2010). Still, our knowledge about the relative contribution of processes operating at different spatial and temporal scales to the tree diversity in Amazonia remains surprisingly limited. Obtaining new scientific insight requires foremost collecting and analyzing additional empirical data. This thesis addresses parts of the existing knowledge gaps by presenting new data of tree inventory plots and by analyzing these in an integrated manner with previously established plots. The thesis thereby aims at deepening our understanding of the tree diversity in Amazonian forests and the ecological mechanisms shaping its current pattern at a local and regional scale. HISTORY OF AMAZONIAN FORESTS Forests have been a permanent feature in Amazonia for the past 55 million years (Ma) (Morley, 2000, Maslin et al., 2005). Throughout its history, Amazonia has experienced dynamic changes in its landscape and climate (Hoorn et al., 2010). Four important historical events have strongly contributed to the current pattern of diversity: a long period of isolation, the Andean uplift, the closing of the Panama isthmus, and changes in global climate (Fig. 1.1) (Burnham and Graham, 1999, Hoorn et al., 2010). This section shortly describes each of these historical events and discusses their impacts on the evolution of tree species diversity in Amazonia. 9 Chapter 1 A D B E C F Figure 1.1: Palaeogeographic maps (A – E) and present configuration of Amazonia (F). A: rainforest diversification in the northern Andes; uplift of the central and northern Andes; a major drainage divide is located in eastern Amazonia; B: intermediately/moderately diverse rainforest in the northern Andes; the onset of the uplift of the northern Andes coincides with a period of a tropical wet climate with an intense dry season in western Amazonia; the major drainage divide is now located in central-eastern Amazonia; C: diverse rainforest in northwestern Amazonia; continued uplift of the central and northern Andes; origin of the Pebas megawetland in western Amazonia; the major drainage divide moves to central-eastern Amazonia; D: diverse rainforest in western Amazonia; diversification of sub-Andean forest; the accelerated uplift of central Andes and northern Andes promotes the establishment of a transcontinental Amazon drainage system; E: diverse rainforest in western Amazonia and disappearance of Amazonian biota in northern Venezuela; the strongly concerted uplift of the entire Andes and a dynamic low-relief river systems of western Amazonia favor the Andean derived nutrients spread over eastern Amazonia; the closure of the Panama isthmus promotes exchange of biota between North and South America (‘Great American Biotic Interchange’ - GABI); F: diverse rainforest, especially in western Amazonia; present configuration of Amazonia. Note that shield areas are geologically stable over the last 55 Ma. Modified after Hoorn and Wesselingh (2010) and Hoorn et al. (2010). 10
Description: