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Pollen Manual and Atlas Amazon Pollen Manual and Atlas Manual e Atlas Palinológico da Amazônia Amazon Pollen Manual and Atlas Manual e Atlas Palinológico da Amazônia Paul Colinvaux Marine Biological Laboratory Woods Hole, Massachusetts, USA formerly at Smithsonian Tropical Research Institute Balboa, Republic of Panama Paulo Eduardo De Oliveira Department of Botany Field Museum of Natural History Chicago, Illinois, USA and Departamento de Paleontologia e Estratigrafia Instituto de Geociências Universidade de São Paulo São Paulo, Brasil and Jorge Enrique Moreno Patiño Smithsonian Tropical Research Institute Balboa, Republic of Panama harwood academic publishers Australia • Canada • China • France • Germany • India • Japan • Luxembourg • Malaysia • The Netherlands • Russia • Singapore • Switzerland This edition published in the Taylor & Francis e-Library, 2005. "To purchase your own copy of this or any of Taylor & Francis or Routledge's collection of thousands of eBooks please go to www.eBookstore.tandf.co.uk." Copyright © 1999 OPA (Overseas Publishers Association) N.V. Published by license under the Harwood Academic Publishers imprint, part of The Gordon and Breach Publishing Group. All rights reserved. No part of this book may be reproduced or utilized in any form or by any means, electronic or mechanical, including photocopying and recording, or by any information storage or retrieval system, without permission in writing from the publisher. Printed in Singapore. Amsteldijk 166 1st Floor 1079 LH Amsterdam The Netherlands British Library Cataloguing in Publication Data Colinvaux, Paul, 1930– Amazon pollen manual and atlas=Manual e atlas palinológico da Amazônia 1. Palynology 2. Pollen, Fossil—Amazon River 3. Lake sediments—Amazon River—Analysis I. Title II. Oliveira, Paulo E.De III. Moreno, Enrique 561.1´3´09811 ISBN 0-203-30517-5 Master e-book ISBN ISBN 0-203-34302-6 (Adobe e-Reader Format) ISBN 90-5702-587-6 (Print Edition) CONTENTS Preface/Prefácio ix 1 Part I/Parte I: Pollen Manual/Manual Palinológico 1 Neotropical Pollen Analysis 3 2 Review of Amazon Paleoecology: The 1997 NWO/Huygens Lecture 17 3 A Guide to Piston Coring of Lake Sediments 32 4 Laboratory Protocols 48 5 Pollen Morphology 65 1 Análise Palinológica Neotropical 69 2 Revisão da Paleoecologia da Amazônia: A aula de NWO/Huygens 85 3 Guia para amostragem, com pistão, de sedimentos lacustres 102 4 Metodologia Palinológica: protocolo e materiais 122 5 Morfologia de Pólen e Esporos 141 Appendix: Sources of Supply 146 Figures/Figuras 149 177 Part II/Parte II: Amazon Pollen Atlas/Atlas Palinológico da Amazônia Introduction/Introdução 179 Key to Major Pollen and Spore Groups 183 Key to Families and Genera by Pollen and Spore Groups 185 Pollen and Spore Descriptions 218 Plates/Pranchas 287 Glossary/Glossário 348 Bibliography 368 Index of Botanical Families 375 Index of Botanical Names 387 About the Authors/Sobre os Autores 398 PREFACE This is both a pollen manual and an atlas. The Manual, Part I, is a practical guide to researching the Amazon past by pollen analysis, with discussions of the theory of tropical pollen analysis followed by “how to” chapters. The Atlas, Part II, is an illustrated guide to pollen found in Amazon sediments. The Manual begins with our assessment of pollen analysis as applied to the Amazon. We find the Amazon pollen record both rich and informative, although its interpretation requires knowledge of the ways in which tropical pollen signals differ from those in temperate latitudes where pollen analysis was invented. We discuss both the interpretation of Amazon pollen data and the use of surface sampling. Chapter 2 reviews the present state of knowledge of the history of Amazon climate and diversity, although much of Amazon history is still controversial or completely unknown. Chapter 3 describes how to core a lake and chapter 4 describes standard laboratory procedures. Coring sediments under the open water of lakes seems forbidding to some who have not yet tried it, but it is in fact a simple exercise in drilling that is easy to learn. We show how the coring is done, what equipment is needed to core, and how to procure the equipment. Likewise, we provide basic laboratory protocols: We begin with subsampling of sediments, then cover pollen extraction, pollen counting, and the presentation of data. For both field and laboratory work, we provide an appendix listing sources of supply, from rubber boats through coring equipment and radiocarbon dating, to exotic pollen markers and software. The final chapter of Part I is a condensed account of pollen and spore morphology, written as a practical guide for the working paleoecologist. The Atlas, which has its own introduction, provides keys, descriptions, and photomicrographs of the pollen taxa most likely encountered in the Amazon rain forest. Our list of 421 pollen taxa seems humble when compared with the more than 80,000 species of vascular plants so far described for the Amazon basin. Yet this list, mostly identified only to family or genus, is sufficient to carry a profound signal of forest communities and climate. The number is far larger than is normally identified in temperate pollen diagrams. Our reduction of the vast Amazon flora to a few hundred identifiable taxa should make Amazonian palynology accessible to the next generation of researchers. A special feature of the Atlas is that the photomicrographs are printed to the scale of one millimeter to one micron, which means that dimensions can be read off directly with an ordinary metric ruler. This Manual and Atlas are the result of many years’ work. De Oliveira and Colinvaux first went coring together in Brazil in 1988, taking advantage of the many years of prior experience of Colinvaux’s Ohio laboratory on lake coring and pollen analysis in Ecuador. Meanwhile, Moreno had compiled a pollen atlas for the tropical forest of Barro Colorado Island, the Smithsonian research site in Panama. When we three were brought together at the Smithsonian Tropical Research Institute for our studies of the environmental history of the Amazon rain forest, the writing of this book was a natural outcome. Many people and organizations contributed to our efforts. The Amazon fieldwork of our laboratories began in Ecuador in association with Drs. Miriam Steinitz-Kannan, Michael C.Miller, Kam-biu Liu, Mark B.Bush, Melanie Riedinger and Mark Schutzow, and with the encouragement of many institutions and people in Ecuador. For the Brazilian fieldwork we thank the Brazilian Research Council, CNPq and the Brazilian Institute for Natural Resources (IBAMA) for permission to collect the Lake Pata sediments, in collaboration with Dr. Elena Franzinelli of the University of Amazonas in Manaus at the Parque National do Pico da Neblina. We thank Dr. Peter Crane of the Field Museum of Natural History in Chicago for his enthusiastic support and provision of a laboratory for the Lake Pata pollen analysis. Our pollen reference collection was built using the herbaria of the Field Museum, the New York Botanical Gardens (with particular help from Dr. Douglas Daly), and the U.S. National Herbarium. Without these great collections, a work of this kind could not be attempted. We thank particularly Dr. Maria Léa Salgado-Labouriau, University of Brasilia, and Dr. Kenitiro Suguio, Institute of Geosciences of the University of São Paulo, for their continued encouragement of our paleoecological studies in Brazil. Mark B.Bush initiated and developed the program of pollen trapping in the Amazon that made possible the naming of many pollen taxa that were up to then unknown. Eduardo Asanza monitored pollen trapping at the Cuyabeno Field Station in Ecuador. Editing and revision of the Portuguese text was by Maria Cleonice L.Sanches. Maria Josefa Villar was invaluable as the trilingual administrative assistant for the project. Salomon Vergara and Marcos Guerra assisted with the production of pollen plates. Eduardo Montenegro worked with artist Alberto Castillo to produce the illustrations for Part I. Our Amazon research has been possible only because of generous support from the U.S. National Science Foundation (NSF), the National Geographic Society, the U.S. National Oceanographic and Atmospheric Administration (NOAA), and the Andrew Mellon Foundation. PREFÁCIO Este livro é, ao mesmo tempo, um manual e um atlas palinológico. O Manual—Parte I— é um guia às técnicas palinológicas adequadas ao estudo do passado da Amazônia e contém discussões sobre a teoria da análise palinológica, seguido de capítulos que ensinam ao leitor os procedimentos básicos envolvidos nessa pesquisa. O Atlas—Parte II—é um guia ilustrado dos tipos polínicos encontrados em sedimentos quaternários da Amazônia. Começamos o Manual com a nossa interpretação da análise palinológica aplicada à Amazônia. O registro palinológico da Região Amazônica é rico e informativo. Entretanto, sua interpretação requer conhecimentos para diferenciar seus sinais polínicos, daqueles encontrados nas regiões de clima temperado, onde foi criada a análise palinológica. Discutimos não só a interpretação dos dados palinológicos da Amazônia, como também o uso de amostragem de sedimentos superficiais. No Capítulo 2 apresentamos uma revisão do estado atual dos conhecimentos da história climática do Quaternário Tardio e da diversidade da Região Amazônica, embora, ainda, muito controverso e desconhecido. No Capítulo 3 descrevemos o procedimento de coleta de sedimentos lacustres e no Capítulo 4, a metodologia palinológica padrão. Embora a coleta de sedimentos em lagos possa parecer
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