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Tree Demography in the Tropical Rain Forest of Guyana

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Tree Demography in the Tropical Rain Forest of Guyana Tree demography in the tropical rain forest of Guyana The Tropenbos-Guyana Programme operates within the framework of the interna­ tional programme of the Tropenbos Foundation. The multi-disciplinary Tropenbos­ Guyana Programme contributes to conservation and wise utilization of forest resourc­ es in Guyana by conducting strategic and applied research and upgrading Guyanese capabilities in the field of forest-relatedsciences. The Tropenbos-Guyana Series publishes results of research projects carried out in the framework of the Tropenbos-Guyana Programme. R.J. Zagt Tree demography in the tropical rain forest of Guyana T ropenbos-Guyana Series 3 Tropenbos-Guyana Programme - Georgetown, Guyana ISBN: 90-393-1741-0 Keywords: population dynamics, tropical rain forest, selective logging © 1997 Tropenbos-Guyana Programme, Roderick Zagt All rights reserved. No part of this publication, apart from bibliographic data and brief quotations in critical reviews, may be reproduced, re-recorded or published in any form including photocopy, microfilm, electronic or electromagnetic record, without written permission. Printed by: Elinkwijk bv Cover: Greenheart forest in inserted in a stamp from former British Guiana, 'Felling Greenheart'. Photo by Roderick Zagt; image-processing by Barr Landman. Tree demography in the tropical rain forest of Guyana De demografie van bomen in het tropische regenwoud van Guyana (met een samenvatting in her Nederlands) Proefschrift ter verkrijging van de graad van doctor aan de Universiteit Utrecht op gezag van de Rector Magnificus, Prof. Dr. H.O. Voorma, ingevolge her besluit van her College van Decanen in her openbaar re verdedigen op woensdag 3 september 1997 des middags te 2.30 uur door Roderick Jan Zagt geboren op 2 mei 1964, te Leiden Promotor: Prof. Dr. M.J.A. Werger Co-promotor: Dr. R.G.A. Boot Verbonden aan de Faculteit Biologie van de Universiteit Utrecht TROPENBOS ��'� � � UniversiteitUtrecht �A� The investigations reported in this thesis were carried out at the Tropenbos-Guyana Programme, Lot 12E Garnerrsrreet, Campbellville, Guyana and the Department of Plant Ecology and Evolutionary Biology, Utrecht University, P.O. Box 800.84, 3508 TB Utrecht, The Netherlands. Contents Chapter I Introduction 7 Chapter 2 Community structure and the demography of primary species 21 in tropical rain forest R.j. Zagt & M.j.A. Werger. Accepted forpttblication in: D.M. Newbery, H.H. T. Prins & ND. Brown. Pop11lation and commttnity dynamics in the tropics. British Ecological SocietySymposittm VolttmeNo 37. Blackwell ScientificP11blishers, Cambridge. Chapter 3 Pre-dispersal and early post-dispersal demography, and 39 reproductive litter production in the tropical tree Dicymbe altsonii in Guyana R.j.Zagt. Accepted forp11blication in journal of Tropical Ecology. Chapter 4 Stem sprouting of Dicymbe altsonii in the tropical rain forest 55 of Guyana: impact of soil type and potential for regeneration R.j. Zagt, E.-j. Malta & M.H. Rijks. Chapter 5 Delayed germination of Chlorocardium rodiei in the tropical 73 rain forest of Guyana R.j.Zagt & D.S. Hammond. Chapter 6 Distribution and demography in relation to light and size for 93 seedlings of three Guyanese tropical rain forest species R.j. Zagt, J.L. Machado & P.B. Reich. Chapter 7 Seedling dynamics of Chlorocardium rodiei in logged-over rain 139 forest in Guyana R.j.Zagt. Chapter 8 The response of tropical trees to logging: a cautious 167 application of matrix models R.j. Zagt & R. G.A. Boot. Chapter 9 Summary and conclusions 215 Samenvatting 223 References 231 Acknowledgements 247 Curriculum Vitae 251 chapter Introduction Over history, western civilisation has built up and maintained an ambiguous rela­ tion with the tropical rain forest. Ever since Fransisco Orellana in 1541 made his in­ voluntary trip over the River Amazon, the Amazonian rain forest has attracted or re­ pelled, but certainly enchanted the western mind. The words tropical rain forest and Amazon evoke images of strange, unseen creatures, riches which abound but often re­ mained elusive, rubber booms and gold rushes, hidden cities and lakes, great endea­ vours and vast failures, opera houses and railroads, stone-age tribes and lost civilisa­ tions, poisoned arrows and plenty of mosquitoes which have all conquered their place in the western mind. Amazonia is a magnifying glass which seems to reject mediocrity. The rivers are wider, the beauty is greater, the heroes more heroic, the scoundrels worse, the forest denser and packed with more species than anywhere else. In these days mediocrity has easy access to Amazonia, but the fascination forits myths remains. Biologists working in the rain forest have not escaped this mystifying force. Scien­ tists turn poets once they set foot into it (examples in Whitmore 1990), and even those scientists who rhyme with numbers find inspiration (Erwin1982, WCMC 1992). The rain forest stands for the most diverse ecosystem on Earth (Connell 1978),.where on 6% of the land area more species are found than in the rest of the world (WCMC 1992), of which 3 species go extinct each hour (Wilson 1989). Keystone species sup­ port extensive and intricate foodwebs (Terborgh 1986). Mast-fruiting is related to black-water rivers Qanzen 1974). Rain forests are growing on sterile soil (cf Brouwer 1996). Seedlings and saplings survive deep shade for up to 30 years (Fanshawe 1947), after which they can still respond to gap opening. Seveney-five percent of all species need gaps to reach the canopy (Hartshorn 1978). Herbivores regulate species richness through distance-dependent mortality Qanzen 1970). Gradually, misconceptions have been corrected, and scientists have increasingly set to quantify these and other aspects of the rain forest, not only in Amazonia. The 1980s and 1990s have become decades of demythologisation of the biological aspects of the rain forest, and although it has lost little of its attraction, at present it is possible to dis­ cuss its enigmas on the base of facts and well-argued theories. The study of the population biology of tropical trees has seen a strong development since the 1960s, when Richards (1952) and Schulz (1960) had just published their pio- 7 chapter neering work. While it is not justified to say that it is now possible to understand the population biology of tropical trees, its study has become a mature ecological under­ taking. There are two important reasons for this emphasis. Biologists have always been intrigued by the enormous diversity of species in tropical rain forests, and the mecha­ nisms that make them coexist in apparently homogeneous areas. Population biology of these species is thought to provide a key to the understanding of this fundamental bio­ logical issue. A second reason for this emphasis is the increasing pressure that tropical rain forests have come to bear from increasing human populations and from economic exploitation, and the realisation that only the management of these forests for timber and other products will be able to diminish this pressure. Sustainable management is only possible with a proper knowledge of the population behaviour of the species to be managed. By far the most significant contribution to the development of the population biolo­ gy of tropical trees since the early days has been the creation and census �f very large permanent sample plots, first and foremost on Barro Colorado Island (Panama), later also in other countries (Condit 1995). These studies have produced an incessant stream of publications, in which new and existing hypotheses were critically assessed using the enormous data-sets that these plots provide. This is a significant improve­ ment, because tropical population biology is an area in biology where traditionally the formulation of hypotheses and paradigms have run far ahead of the field work and ex­ periments to test them Qanzen 1970, Connell 1971, Hartshorn 1978, Hubbell 1979). Still, what these studies gain in scale, they loose in detail, and therefore other, smaller studies, however sometimes encumbered by the consequences of the same species rich­ ness that they study (small sample sizes, taxonomic uncertainty), provide invaluable detail to the knowledge generated in the large plots. This dissertation is based on the results of a (comparably) small population study of three tree species from the rain forest of Guyana. It describes the population dynamics of three species differingin regeneration strategy, and links this to timber exploitation. While the objectives quite specifically concentrate on assessing the consequences of ex­ ploitation on the population dynamics of these species, a large part of the dissertation is spent on quantifying and understanding specific components of the life cycle of these species, and on reviewing their population dynamics in the light of the main hy­ potheses that are put forward to explain species coexistence and succession in tropical rain forests. A fully documented introduction of hypotheses concerning species coexistence in tropical rain forests, and an identification of parameters that are thought to shape the course of succession in rain forests is provided in Chapter 2 of this thesis. The ideas forwarded in this Chapter are the fundament of much of the work reported elsewhere in the thesis. The following introduction is, therefore, limited to a clarification of some concepts and choices that have shaped this thesis, and an introduction to Guyana and the Tro­ penbos Programme that have coloured it. 8 chapter Population biology and tropical trees Demography is the study of the quantitative aspects of populations; the population biology investigates the factors that govern these aspects (Harper 1979). The number of individuals in a plant population changes as a result of birth and death, and in the seed stage by immigration and emigration. Between birth and death plants grow and only if sufficient individuals grow enough to reach the reproductive stage, the popula­ tion is perpetuated. Hence, the population dynamics of the species in this study are studied by measuring and interpreting their rates of growth, survival and reproduction. Nature offersmany ways to grow to maturity.
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