Ecological Studies Analysis and Synthesis Edited by WD. Billings, Durham (USA) F. Golley, Athens (USA) O.L. Lange, Wtirzburg (FRG) 1.S. Olson, Oak Ridge (USA) H. Remmert, Marburg (FRG) Volume 63 Ecological Studies Volume 52 Volume 58 Air Pollution by Photochemical Oxidants Ecology of Biological Invasions of North Formation, Transport, Control and America and Hawaii Effects on Plants Edited by H.A. Mooney and J.A. Drake Edited by Robert Guderian 1986. X. 320p., 25 figures. cloth 1985. XI, 346p., 54 figures. cloth ISBN 0-387-96289-1 ISBN 0-387-13966-4 Volume 59 Volume 53 Acid Deposition and the Acidification The Gavish Sabkha of Soils and Waters A Model of a Hypersaline Ecosystem An Analysis Edited by G.M. Friedman and W. E. By 1.0. Reuss and D.W. Johnson Krumbein 1986. VIII. 120p., 37 figures. cloth 1985. X. 484p., 246 figures. cloth ISBN 0-387-96290-5 ISBN 0-387-15245-8 Volume 60 Volume 54 Amazonian Rain Forests Tidal Flat Ecology Ecosystem Disturbance and Recovery An Experimental Approach to Species Edited by Carl F. Jordan Interactions 1987. X. 133p .. 55 figures. cloth By Karsten Reise ISBN 0-387-96397-9 1985. X. 198p .. 69 figures. cloth ISBN 0-387-15447-7 Volume 61 Potentials and Limitations Volume 55 of Ecosystem Analysis A Eutrophic Lake Edited by E.-D. Schulze and H. Zwiilfer Lake Mendota, Wisconsin 1987. XII. 435p .. 141 figures. cloth By Thomas D. Brock ISBN 0-387-17138-X 1985. XII. 308p .. 82 figures. cloth ISBN 0-387-96184-4 Volume 62 Volume 56 Frost Survival of Plants Resources and Society By A. Sakai and W. Larcher A Systems Ecology Study of the Island 1987. XI, 321 p .. 200 figures. n tables. cloth of Gotland, Sweden ISBN 0-387-17332-3 By James J. Zucchetto and Ann-Mari Jansson Volume 63 1985. X. 248p .. 70 figures. cloth Long-Term Forest Dynamics ISBN 0-387-96151-8 of the Temperate Zone By Paul A. Delcourt and Hazel R. Delcourt Volume 57 1987. XIV. 450p .. 90 figures. 333 maps. cloth Forest Ecosystems in the Alaskan Taiga ISBN 0-387-96495-9 A Synthesis of Structure and Function Edited by K. Van Cleve. F.S. Chapin III. Volume 64 L.A. Viereck. C.T. Dyrness and Landscape Heterogeneity and Disturbance P.W. Flanagan Edited by Monica Goigel Turner 1986. X, 240p .. 81 figures. cloth 1987. XI, 241 p .. 56 figures. cloth ISBN 0-387-96251-4 ISBN 0-387-96497-5 Paul A. Delcourt Hazel R. Delcourt Long-Term Forest Dynamics of the Temperate Zone A Case Study of Late-Quaternary Forests in Eastern North America With 90 Illustrations in 131 Parts and 333 Maps Springer-Verlag New York Berlin Heidelberg London Paris Tokyo PAUL A. DELCOURT HAZEL R. DELCOURT Program for Quaternary Studies Program for Quaternary Studies of the Southeastern United States of the Southeastern United States Graduate Program in Ecology Graduate Program in Ecology Department of Geological Sciences Department of Botany University of Tennessee University of Tennessee Knoxville. TN 37996. USA Knoxville. TN 37996. USA Library of Congress Cataloging-in-Publication Data Delcourt. Paul A. Long-term forest dynamics of the Temperate Zone. (Ecological studies; v. 63) Bibliography: p. Includes index. I. Pollen. Fossil. 2. Paleoecology-North America. 3. Paleobotany-Quaternary. I. Delcourt. Hazel R. II. Title. III. Series. QE993.2.D45 1987 561' \3 87-12859 ISBN-13: 978-1-4612-9136-7 e-ISBN-13: 978-1-4612-4740-1 DOl: lO.lO07/978-1-4612-4740-1 © 1987 by Springer-Verlag New York Inc. Softcover reprint of the hardcover 1st edition 1987 All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer-Verlag. 175 Fifth Avenue. New York. New York 10010. USA). except for brief excerpts in connection with reviews or scholarly analysis. Use in connection with any form of information storage and retrieval. electronic adaptation. computer software. or by similar or dissimilar meth odology now known or hereafter developed is forbidden. The use of general descriptive names. trade names. trademarks. etc. in this publication. even if the former are not especially identified. is not to be taken as a sign that such names. as understood by the Trade Marks and Merchandise Marks Act, may accordingly be used freely by anyone. Typeset by David Seham Associates. Inc .. Metuchen. New Jersey. 987654321 ISBN-13: 978-1-4612-9136-7 Springer-Verlag New York Berlin Heidelberg To our Paleoecological Mentors Ronald O. Kapp Herbert E. Wright, Jr. Preface The synthesis presented in this volume is a direct outgrowth of our ten-year FORMAP Project (Forest Mapping Across Eastern North America from 20,000 yr B.P. to the Present). Many previous research efforts in paleoecology have used plant-fossil evidence as proxy information for primarily geologic or climatic reconstructions or as a bio stratigraphic basis for correlation of regional events. In contrast, in this book, we deal with ecological questions that require a holistic perspective that integrates the interactions of biota with their dynamically changing environments over time scales up to tens of thousands of years. In the FORMAP Project, our major research objective has been to use late-Quaternary plant-ecological data sets to evaluate long-term patterns and processes in forest de velopment. In order to accomplish this objective, we have prepared subcontinent-scale calibrations that quantitatively relate the production and dispersal of arboreal pollen to dominance in the vegetation for the major tree types of eastern North America. Quantification of pollen-vegetation relationships provides a basis for developing quan titative plant-ecological data sets that allow further ecological analysis of both individual taxa and forest communities through time. Application of these calibrations to fossil pollen records for interpreting forest history thus represents a fundamental step beyond traditional summaries based upon pollen percentages. The scope of this book includes an introduction to the nature of long-term vegetational change, dealing with different kinds of environmental and biotic changes that result in dynamic patterns and processes in forest communities. Quantification of modem pollen-vegetation relationships permits the development of techniques for reconstruction viii Preface of past forests, when applied to plant-fossil sequences in, for example, a case study of eastern North America. We use these pollen-vegetation calibrations to reconstruct forest history based upon radiocarbon-dated, late-Quaternary fossil-pollen sequences of 162 sites distributed across the eastem half of North America. The resulting FORMAP time series of maps depict the changing distribution and dominance patterns for pop ulations of major temperate and boreal tree taxa. Areographic analysis of the map patterns provides new insights into migrational strategies of trees during times of en vironmental change. The role of competition in structuring past forest communities is examined using multiple-taxa population-growth models in conjunction with empirical paleoecological data. Ordination of the paleoecological data using Detrended Corre spondence Analysis gives information concerning changes in position and steepness of ecotones. This study represents the first use of DCA (DECORANA) in direct gradient analysis of changing ecoclines through both space and time on a scale relevant to the dynamic development of temperate forest communities. We also develop perspectives in Quaternary landscape ecology, integrating climatic and geomorphic processes with changes in vegetational patterns through time. We compare late-Quaternary pattern and process in development of eastern North American forests with forest dynamics of the Temperate Zone of Europe. The Appendix to this volume includes a listing with supplementary information for all fossil-pollen sites used in the FORMAP project. The complete data bases for modem forest composition and pollen spectra, as well as all fossil-pollen data and forest re constructions, are on file at the Program for Quaternary Studies of the Southeastern United States, University of Tennessee, Knoxville. We acknowledge financial support from the Ecology Program of the United States National Science Foundation for the FORMAP Project (grants DEB-80-04168 and BSR-83-06915) and for related research (grants BSR-83-00345 and BSR-84-15652). We are grateful to P. Wright and D. Broach for their assistance in computer program ming. We express our appreciation to the many palynologists who have graciously made their late-Quatemary pollen data available to us for this purpose. We collaborated with T. Webb III in the compilation and exchange of both modern and fossil pollen data for eastern North America. We thank I.e. Prentice for his advice concerning appropriate statistical techniques for evaluating paleoecological data. We thank F. B. Golley, S.A. Hall, and P.S. White for their constructive reviews of earlier drafts of this manuscript. This volume constitutes Contribution No. 43 of the Program for Qua ternary Studies of the Southeastern United States, University of Tennessee, Knox ville. This book also represents a United States Contribution to the International Geological Correlation Programme O. G. e. P.) Project 158B, dealing with late-Quaternary biotic and environmental changes in the Temperate Zone. Knoxville, Tennessee, 1987 Paul A. Delcourt Hazel R. Delcourt Contents Preface vii 1. The Nature of Long-Term Vegetational Change Long-term patterns and processes in forest communities Spatial and temporal scale 3 The scale paradigm 3 The mega-scale domain 4 The macro-scale domain 6 The micro-scale domain 8 Quaternary environmental changes as forcing functions 9 Episodes of continental glaciation in Earth history 9 Pacemaker of the late-Cenozoic Ice Ages 10 Leads and lags in response of environment and biota to climatic change 13 Anthropogenic influences 16 Models of stability and dynamics of forest communities 19 A north-to-south transect through the last 20,000 years 19 The role of changing disturbance regimes 23 Conclusions 28 x Contents 2. Modern Pollen-Vegetation Relationships 29 Production and dispersal of pollen grains 29 Pollen production and mechanisms for dispersal 30 Temporal patterns of pollen dispersal 32 Spatial patterns of pollen dispersal 36 Calibration of the relationship of modern pollen assemblages to extant forests 42 The representation of temperate and boreal tree taxa in modern pollen samples and forest vegetation 46 Regional pollen-vegetation relationships in eastern North America 47 Regional studies elsewhere in the northern Temperate Zone 49 Subcontinent-scale pollen-vegetation calibrations in eastern North America 50 Conclusions 60 3. Reconstructing Long-Term Forest Changes From Fossil-Pollen Data 62 Comparison of quantitative techniques for reconstruction of past vegetation 62 Taxon calibrations 63 Modern analogue methods 64 Forest simulation models 67 A case study in comparison of taxon calibrations, modern analogue techniques, and simulation models as different approaches for quantifying past vegetation 71 The late-Quaternary pollen record from Anderson Pond 71 Results of taxon calibrations applied to Anderson Pond 71 Results of analogue measures applied to Anderson Pond 77 Results of the FORET simulation model at Anderson Pond 81 Advantages and disadvantages of quantitative methods for forest reconstruction 82 Generating late-Quaternary plant-ecological data sets 83 Conclusions 84 4. Vegetation Map Patterns at the Biome Level 85 Constructing paleogeographic maps 85 Physical geography 85 Site selection and chronologies 86 Paleovegetation maps for eastern North America: 20,000 yr B.P. to the present 88 Characterization of modern vegetation by pollen assemblages 89 Contents xi Selecting time planes for mapping 94 Interpretation of paleovegetation maps 95 Patterns of paleoclimate inferred from maps of paleovegetation 99 The relationship of modern climatic regions to vegetation 99 Biotic responses to changing atmospheric circulation patterns during the late Quaternary 100 Conclusions 106 5. Tree Population Dynamics During the Past 20,000 Years 107 Population trajectories through time 108 Documentation of changing number and density of paleoecological sites through time 108 Minimum, maximum, and mean values of dominance through time 109 Determination of contour intervals of dominance scaled to the population characteristics of each taxon 110 Temporal changes in areal extent and dominance structure of tree populations 110 Contoured paleo-dominance maps for major tree taxa in eastern North America III Fir (Abies) II I Maple (Acer) 122 Birch (Betula) 131 Hickory (Carya) 142 Hackberry (Celtis) 151 Cedars and cypress (Cupressaceae and Taxodiaceae) 158 Beech (Fagus grandifolia) 169 Ash (Fraxinus) 179 Walnut (Juglans) 188 Tamarack (Larix) 195 Tupelo (Nyssa) 205 Spruce (Picea) 213 Pine (Pinus) 224 Aspen or Cottonwood (Populus) 236 Oak (Quercus) 246 Willow (Salix) 256 Basswood (Tilia) 264 Hemlock (Tsuga) 272 Elm (Ulmus) 282 Conclusions 291 6. Late-Quaternary Migrational Strategies of Tree Species 292 Ecological areography 293 Measurements of perimeter/V area 294 Ecological areography of eastern North American tree taxa 295