THE GEOLOGICAL RECORD OF ECOLOGICAL DYNAMICS

UNDERSTANDING THE BIOTIC EFFECTS OF FUTURE ENVIRONMENTAL CHANGE

Committee on the Geologic Record of Biosphere Dynamics

Board on Earth Sciences and Resources

Board on Life Sciences

Division on Earth and Life Studies

NATIONAL RESEARCH COUNCIL OF THE NATIONAL ACADEMIES

THE NATIONAL ACADEMIES PRESS
Washington, D.C. www.nap.edu



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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change THE GEOLOGICAL RECORD OF ECOLOGICAL DYNAMICS UNDERSTANDING THE BIOTIC EFFECTS OF FUTURE ENVIRONMENTAL CHANGE Committee on the Geologic Record of Biosphere Dynamics Board on Earth Sciences and Resources Board on Life Sciences Division on Earth and Life Studies NATIONAL RESEARCH COUNCIL OF THE NATIONAL ACADEMIES THE NATIONAL ACADEMIES PRESS Washington, D.C. www.nap.edu

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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change THE NATIONAL ACADEMIES PRESS 500 Fifth Street, N.W. Washington, DC 20001 NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research Council, whose members are drawn from the councils of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The members of the committee responsible for the report were chosen for their special competences and with regard for appropriate balance. The opinions, findings, and conclusions or recommendations contained in this document are those of the authors and do not necessarily reflect the views of the National Science Foundation or the U.S. Geological Survey. Mention of trade names or commercial products does not constitute their endorsement by the U.S. Government. Supported by the National Science Foundation (NSF) under Contract No. 0106060 and by the U.S. Geological Survey (USGS), Department of the Interior, under Award No. 01HQAG0216. International Standard Book Number 0-309-09580-8 (Book) International Standard Book Number 0-309-54844-6 (PDF) Library of Congress Control Number: 2005924162 Additional copies of this report are available from the National Academies Press, 500 Fifth Street, N.W., Lockbox 285, Washington, DC 20055; (800) 624-6242 or (202) 334-3313 (in the Washington metropolitan area); Internet http://www.nap.edu Cover: Background photograph shows mollusk shells on the Colorado River delta in Mexico, which record the biotas and environmental conditions prior to upstream diversion of freshwater. Radiocarbon, amino acid, and growth line analyses of the shells, together with oxygen and carbon stable isotope analyses, provide detailed chronologies of temperature, salinity, productivity, and other environmental parameters. Photo courtesy Karl W. Flessa. Outcrop photograph shows rhythmically bedded Late Cretaceous (~87 Ma) limestones and shales in the Terlingua Creek canyon of West Texas. The geologic record can be used as an ecological laboratory when rocks preserve a near-continuous sequence of repeated environmental and biotic change; photo courtesy Brad Sageman. Copyright 2005 by the National Academy of Sciences. All rights reserved. Printed in the United States of America.

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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change THE NATIONAL ACADEMIES Advisers to the Nation on Science, Engineering, and Medicine The National Academy of Sciences is a private, nonprofit, self-perpetuating society of distinguished scholars engaged in scientific and engineering research, dedicated to the furtherance of science and technology and to their use for the general welfare. Upon the authority of the charter granted to it by the Congress in 1863, the Academy has a mandate that requires it to advise the federal government on scientific and technical matters. Dr. Bruce M. Alberts is president of the National Academy of Sciences. The National Academy of Engineering was established in 1964, under the charter of the National Academy of Sciences, as a parallel organization of outstanding engineers. It is autonomous in its administration and in the selection of its members, sharing with the National Academy of Sciences the responsibility for advising the federal government. The National Academy of Engineering also sponsors engineering programs aimed at meeting national needs, encourages education and research, and recognizes the superior achievements of engineers. Dr. Wm. A. Wulf is president of the National Academy of Engineering. The Institute of Medicine was established in 1970 by the National Academy of Sciences to secure the services of eminent members of appropriate professions in the examination of policy matters pertaining to the health of the public. The Institute acts under the responsibility given to the National Academy of Sciences by its congressional charter to be an adviser to the federal government and, upon its own initiative, to identify issues of medical care, research, and education. Dr. Harvey V. Fineberg is president of the Institute of Medicine. The National Research Council was organized by the National Academy of Sciences in 1916 to associate the broad community of science and technology with the Academy’s purposes of furthering knowledge and advising the federal government. Functioning in accordance with general policies determined by the Academy, the Council has become the principal operating agency of both the National Academy of Sciences and the National Academy of Engineering in providing services to the government, the public, and the scientific and engineering communities. The Council is administered jointly by both Academies and the Institute of Medicine. Dr. Bruce M. Alberts and Dr. Wm. A. Wulf are chair and vice chair, respectively, of the National Research Council. www.national-academies.org

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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change COMMITTEE ON THE GEOLOGIC RECORD OF BIOSPHERE DYNAMICS: THE KEY TO UNDERSTANDING THE BIOTIC EFFECTS OF FUTURE ENVIRONMENTAL CHANGE KARL W. FLESSA, Chair, University of Arizona, Tucson STEPHEN T. JACKSON, Vice Chair, University of Wyoming, Laramie JOHN D. ABER, University of New Hampshire, Durham MICHAEL A. ARTHUR, Pennsylvania State University, University Park PETER R. CRANE, Royal Botanic Gardens, Kew, United Kingdom DOUGLAS H. ERWIN, Smithsonian Institution, National Museum of Natural History, Washington, D.C. RUSSELL W. GRAHAM, Pennsylvania State University, University Park JEREMY B.C. JACKSON, Scripps Institution of Oceanography, University of California, San Diego SUSAN M. KIDWELL, The University of Chicago, Illinois CHRISTOPHER G. MAPLES, Desert Research Institute, Las Vegas and Reno, Nevada CHARLES H. PETERSON, University of North Carolina, Morehead City O. JAMES REICHMAN, University of California, Santa Barbara Liaison from the Board on Earth Sciences and Resources DAVID L. DILCHER, University of Florida, Gainesville National Research Council Staff DAVID A. FEARY, Study Director (Board on Earth Sciences and Resources) ROBIN A. SCHOEN, Senior Program Officer (Board on Life Sciences) JENNIFER T. ESTEP, Administrative Associate RADHIKA S. CHARI, Senior Project Assistant (until 04/04) AMANDA M. ROBERTS, Project Assistant (from 06/04)

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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change BOARD ON EARTH SCIENCES AND RESOURCES GEORGE M. HORNBERGER, Chair, University of Virginia, Charlottesville M. LEE ALLISON, Kansas Geological Survey, Lawrence JILL BANFIELD, University of California, Berkeley STEVEN R. BOHLEN, Joint Oceanographic Institutions, Washington, D.C. ADAM M. DZIEWONSKI, Harvard University, Cambridge, Massachusetts RHEA L. GRAHAM, New Mexico Interstate Stream Commission, Albuquerque ROBYN HANNIGAN, Arkansas State University, Jonesboro V. RAMA MURTHY, University of Minnesota, Minneapolis RAYMOND A. PRICE, Queen’s University, Ontario, Canada MARK SCHAEFER, NatureServe, Arlington, Virginia STEVEN M. STANLEY, Johns Hopkins University, Baltimore, Maryland BILLIE L. TURNER II, Clark University, Worcester, Massachusetts STEPHEN G. WELLS, Desert Research Institute, Reno, Nevada THOMAS J. WILBANKS, Oak Ridge National Laboratory, Tennessee National Research Council Staff ANTHONY R. DE SOUZA, Director TAMARA L. DICKINSON, Senior Program Officer DAVID A. FEARY, Senior Program Officer ANNE M. LINN, Senior Program Officer ANN G. FRAZIER, Program Officer RONALD F. ABLER, Senior Scholar TANJA E. PILZAK, Research Associate CAETLIN M. OFIESH, Research Assistant JENNIFER T. ESTEP, Administrative Associate VERNA J. BOWEN, Administrative Assistant MARIA A. ALEJANDRO, Program Assistant JAMES B. DAVIS, Program Assistant AMANDA M. ROBERTS, Program Assistant

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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change BOARD ON LIFE SCIENCES COREY S. GOODMAN, Chair, University of California, Berkeley RUTH L. BERKELMAN, Emory University, Atlanta, Georgia R. ALTA CHARO, University of Wisconsin, Madison DENNIS CHOI, Merck Research Laboratories, West Point, Pennsylvania JOANNE CHORY, The Salk Institute for Biological Studies, La Jolla, California JEFFREY L. DANGL, University of North Carolina, Chapel Hill PAUL R. ERLICH, Stanford University, Palo Alto, California JAMES M. GENTILE, Hope College, Holland, Michigan LINDA E. GREER, Natural Resources Defense Council, Washington, D.C. ED HARLOW, Harvard Medical School, Cambridge, Massachusetts DAVID M. HILLIS, University of Texas, Austin KENNETH H. KELLER, University of Minnesota, Minneapolis RANDALL MURCH, Institute for Defense Analyses, Alexandria, Virginia GREGORY A. PETSKO, Brandeis University, Waltham, Massachusetts STUART L. PIMM, Duke University, Durham, North Carolina BARBARA A. SCHAAL, Washington University, St. Louis, Missouri JAMES M. TIEDJE, Michigan State University, East Lansing KEITH R. YAMAMOTO, University of California, San Francisco National Research Council Staff FRANCES E. SHARPLES, Director ROBIN A. SCHOEN, Senior Program Officer ROBERT T. YUAN, Senior Program Officer KERRY A. BRENNER, Program Officer MARILEE K. SHELTON-DAVENPORT, Program Officer EVONNE P.Y. TANG, Program Officer ADAM P. FAGEN, Postdoctoral Fellow BRENDAN BRADLEY, Program Assistant/Research Intern MILTON MULDROW, Program Assistant SETH STRONGIN, Program Assistant DENISE GROSSHANS, Financial Associate

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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change Preface The economic and environmental security of the United States and the world depends on the ecosystem services and functions provided by natural and managed biotic systems. It is vital therefore that we learn how these systems respond to both natural and anthropogenic change. Predicting and managing the biotic response to future environmental change require a sound scientific understanding of the rate and character of past biotic responses to environmental perturbations. The converse also applies—biotic systems, through their effects on the characteristics of the earth’s surface and on biogeochemical cycles, affect the climate system. Understanding this feedback is a crucial requirement for predicting both future climate and the response of the biosphere to climate change. The geologic record is vital for understanding the complex interactions between environmental and biotic change; the last 200 years of direct scientific observations is too short a time period to allow direct observation of the range of environmental conditions we are likely to encounter in the future, too short to expose the nature of long-term ecological processes, and too short to fully reveal the ecological legacy of past events. The geologic record also provides critical insight into the character of biotic systems unaffected by human activities. Recent technological and conceptual advances in the earth and life sciences make analyses of the geologic record of past biotic change especially powerful and relevant. Ecological studies are now revealing the importance of past events, long-term environmental change, and evolutionary processes in the structure and function of living ecosystems. Significant improvements in dating geological materials and correlating

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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change events among sites now allow the resolution of rates of environmental and biotic change, as well as the identification of short-term events and their effects over large areas. Two decades of intense research on the processes of fossilization and on analytical methods to detect and compensate for incomplete and biased records now allow scientists to discern true ecologic patterns in fossil assemblages and true evolutionary signals in the fossil record. The development of precise biogeochemical indicators of past environmental and biotic conditions now permits high-resolution reconstructions of past biotas and their variability. The power of these developments has been greatly increased by the development of large databases and tools for data dissemination, integration, and visualization. Such databases and database tools provide the foundation for regional and global syntheses to assess the ecological and evolutionary impacts of environmental changes across a range of timescales. These advances set the stage for collaborative efforts among earth scientists and life scientists for innovative and integrative examinations of the geological record of past environmental and biotic change. In light of these advances—and the potential for collaborative, interdisciplinary research on the nature of biotic response to future environmental change—the National Science Foundation and the U.S. Geological Survey requested that the National Academies assess the scientific opportunities provided by the geologic record of biosphere dynamics, and recommend how academic and agency scientists could best exploit those opportunities for the nation’s benefit. The committee, composed of both life scientists and earth scientists, met three times as a group to review existing agency and academic research efforts, to hear testimony solicited from expert scientists, to formulate its recommendations, and to write key components of the report. The concept of biosphere dynamics encompasses any and all changes in the earth system that involve living organisms, regardless of temporal or spatial scale. Environmental changes, including climate change, tectonic activity, and sea level changes, have driven evolutionary and ecological dynamics at timescales ranging from thousands to hundreds of millions of years. Evolutionary novelties, ranging from the origin of oxygenic photosynthesis to the origin of our own species, have led to a broad range of environmental and ecological responses. Faced with the vast array of phenomena that could be subsumed under the term “biosphere dynamics,” and recognizing the basic scientific and applied resource management issues facing society as local and global environmental change accelerates, the committee chose to concentrate its efforts on the geologic record of “ecological dynamics”—the changes and interactions in the earth-life system expressed as alterations in such features as species distributions, species abundance, environment, and climate. While recognizing the sig-

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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change nificance of evolutionary responses to environmental change throughout the history of life, the committee felt that a study that also treated evolutionary dynamics in great detail would require resources and expertise far beyond those available to it. This report is the result of many additional hours of discussions, writing, reviewing, and editing by a committee whose breadth and expertise was essential to the quality of the final product. We are also grateful for David Feary’s skillful guidance and editorial skills and for the support of National Research Council officials and staff. Karl Flessa Chair Stephen Jackson Vice Chair

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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change Acknowledgments This report was greatly enhanced by the participants in the two public committee meetings held as part of this study. The committee would first like to acknowledge the efforts of those who gave presentations at meetings: Enriqueta Barrera, David Blockstein, Lynn Brewster-Wingard, Scott Collins, Rachael Craig, Bruce Fouke, Susan Haseltine, Lars Hedin, Patrick Leahy, Brian Maurer, Michael Meyer, Donald Rice, Craig Schiffries, Steven Stanley, David Verardo, Debra Willard, and Herman Zimmerman. These talks helped set the stage for fruitful discussions in the sessions that followed. The committee is also grateful to Jill Banfield, Tim Baumgartner, Julio L. Betancourt, Cinzia Cervato, David Jablonski, Dennis H. Knight, Peter Leavitt, Thomas Olszewski, William A. Reiners, Brad Sageman, Bryan Shuman, and John W. Williams, who provided important information, commentary, and material for this report. We also thank Dena Smith and the University of Colorado Museum for providing meeting facilities. This report has been reviewed in draft form by individuals chosen for their diverse perspectives and technical expertise, in accordance with procedures approved by the National Resarch Council’s Report Review Committee. The purpose of this independent review is to provide candid and critical comments that will assist the institution in making its published report as sound as possible and to ensure that the report meets institutional standards for objectivity, evidence, and responsiveness to the study charge. The review comments and draft manuscript remain confidential to protect the integrity of the deliberative process. We wish to thank the following individuals for their participation in the review of this report:

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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change James H. Brown, Department of Biology, University of New Mexico, Albuquerque Margaret B. Davis, Department of Ecology, Evolution and Behavior (emeritus), University of Minnesota Andrew H. Knoll, Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, Massachusetts Charles R. Marshall, Department of Earth and Planetary Sciences, Harvard University, Cambridge, Massachusetts Steven M. Stanley, Morton K. Blaustein Department of Earth and Planetary Sciences, Johns Hopkins University, Baltimore, Maryland Roger E. Summons, Department of Earth, Atmospheric, and Planetary Sciences, Massachusetts Institute of Technology, Cambridge Peter M. Vitousek, Department of Biological Sciences, Stanford University, California Although the reviewers listed above have provided many constructive comments and suggestions, they were not asked to endorse the conclusions or recommendations nor did they see the final draft of the report before its release. The review of this report was overseen by P. Geoffrey Feiss, College of William and Mary, Williamsburg, Virginia, appointed by the Divison on Earth and Life Studies, who was responsible for making certain that an independent examination of this report was carried out in accordance with institutional procedures and that all review comments were carefully considered. Responsibility for the final content of this report rests entirely with the authoring committee and the institution.

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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change Contents     EXECUTIVE SUMMARY   1 1   INTRODUCTION   12      Background to This Report,   15      Environmental Sciences and the Geologic Record of Ecological Dynamics,   17      Understanding Processes—Different Perceptions,   19      Status of Research Support,   20      Committee Charge and Scope of the Study,   28 2   THE NATURE AND VALUE OF GEOHISTORICAL INFORMATION   30      The Rationale of Geohistorical Analysis,   30      Completeness of the Geologic Record,   33      Inferring Environmental Conditions—Resolution and Confidence,   37      Biological Insights Contained in the Geologic Record,   44      Precision and Accuracy of Geohistorical Records,   62      Impact of Database Technologies and Synthetic Studies,   71      Summary and Outlook—Strategies for Geohistorical Analysis,   73 3   RESEARCH OPPORTUNITIES   75      The Geologic Record as an Ecological Laboratory,   77      Ecological Responses to Past Climate Change,   98      Ecological Legacies of Societal Activities,   117

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The Geological Record of Ecological Dynamics: Understanding the Biotic Effects of Future Environmental Change 4   THE CULTURE OF COLLABORATIVE RESEARCH   131      Promoting the Culture of Collaboration,   133      Opportunities for Collaboration,   135      Summary,   147 5   CONCLUSIONS AND RECOMMENDATIONS   148      Initiative 1: The Geologic Record as an Ecological Laboratory,   150      Initiative 2: Ecological Responses to Past Climate Change,   151      Initiative 3: Ecological Legacies of Societal Activities,   152      Facilities and Infrastructure,   153      Implementation,   162     REFERENCES   165     APPENDIXES         A  Committee and Staff Biographies   193     B  Acronyms and Abbreviations   199