The National Academies

Advisers to the Nation on Science, Engineering\ and Medicine

In the Light of Evolution Volume II: Biodiversity and Extinction

In the Light of Evolution Volume II: Biodiversity and Extinction




THE NATIONAL ACADEMIES PRESS 500 Fifth Street., N.W. Washington, DC 20001

This volume is based on the Arthur M. Sackler Colloquium of the National Academy of Sciences, "In the Light of Evolution II: Biodiversity and Extinction," held December 7-8, 2007, at the Arnold and Mabel Beckman Center of the National Academies of Sciences and Engineering in Irvine, California. The articles appearing in these pages were contributed by speakers at the colloquium and have been anonymously reviewed. Any opinions, findings, conclusions, or recommendations expressed in this volume are those of the authors and do not necessarily reflect the view of the National Academy of Sciences.

In the light of evolution / John C. Avise and Francisco J. Ayala, editors. p. cm.

Vol. I based on a colloquium of the National Academy of Sciences, held December 1-2, 2006, in Irvine, California. Includes bibliographical references. ISBN-13: 978-0-309-10405-0 (hardcover) ISBN-10: 0-309-10405-X (hardcover) ISBN-13: 978-0-309-66786-9 (pdf) ISBN-10: 0-309-66786-0 (pdf)

1. Evolution (Biology)—Congresses. I. Avise, John C, 1948-. II. Ayala, Francisco José, 1934-III. National Academy of Sciences (U.S.) QH359.I55 2007 576.8--dc22


Additional copies of this book are available from the National Academies Press, 500 Fifth St., N.W., Lockbox 285, Washington, DC 10055; (800) 624-6242 or (202) 334-3313 (in the Washington metropolitan area); Internet, http: //

Cover image: Stylized turtle, street art from Quito, Ecuador (artist unknown). Driven by human agriculture and industry, the Earth is currently undergoing the sixth mass extinction episode in its history. This image symbolizes both human industry and the countless unknown species that are disappearing before they can be catalogued and studied. Image courtesy of John C. Avise.

Copyright 2008 by the National Academy of Sciences. All rights reserved. Printed in the United States of America


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. Ralph J. Cicerone 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. Charles M. Vest 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. Ralph J. Cicerone and Dr. Charles M. Vest are chair and vice chair, respectively, of the National Research Council.

Arthur M. Sackler, M.D. 1913-1987

Born in Brooklyn, New York, Arthur M. Sackler was educated in the arts, sciences, and humanities at New York University. These interests remained the focus of his life, as he became widely known as a scientist, art collector, and philanthropist, endowing institutions of learning and culture throughout the world.

He felt that his fundamental role was as a doctor, a vocation he decided upon at the age of four. After completing his internship and service as house physician at Lincoln Hospital in New York City, he became a resident in psychiatry at Creedmoor State Hospital. There, in the 1940s, he started research that resulted in more than 150 papers in neuroendocri-nology, psychiatry, and experimental medicine. He considered his scientific research in the metabolic basis of schizophrenia his most significant contribution to science and served as editor of the Journal of Clinical and Experimental Psychobiology from 1950 to 1962. In 1960 he started publication of Medical Tribune, a weekly medical newspaper that reached over one million readers in 20 countries. He established the Laboratories for Therapeutic Research in 1938, a facility in New York for basic research that he directed until 1983.

As a generous benefactor to the causes of medicine and basic science, Arthur Sackler built and contributed to a wide range of scientific institutions: the Sackler School of Medicine established in 1972 at Tel Aviv University, Tel Aviv, Israel; the Sackler Institute of Graduate Biomedical Science at New York University, founded in 1980; the Arthur M. Sackler Science Center dedicated in 1985 at Clark University, Worcester, Massachusetts; and the Sackler School of Graduate Biomedical Sciences, established in 1980, and the Arthur M. Sackler Center for Health Communications, established in 1986, both at Tufts University, Boston, Massachusetts.

His pre-eminence in the art world is already legendary. According to his wife Jillian, one of his favorite relaxations was to visit museums and art galleries and pick out great pieces others had overlooked. His interest in art is reflected in his philanthropy; he endowed galleries at the Metropolitan Museum of Art and Princeton University, a museum at Harvard vii

University, and the Arthur M. Sackler Gallery of Asian Art in Washington, D.C. True to his oft-stated determination to create bridges between peoples, he offered to build a teaching museum in China, which Jillian made possible after his death, and in 1993 opened the Arthur M. Sackler Museum of Art and Archaeology at Peking University in Beijing.

In a world that often sees science and art as two separate cultures, Arthur Sackler saw them as inextricably related. In a speech given at the State University of New York at Stony Brook, Some reflections on the arts, sciences and humanities, a year before his death, he observed: ''Communication is, for me, the primum movens of all culture. In the arts . . . I find the emotional component most moving. In science, it is the intellectual content. Both are deeply interlinked in the humanities." The Arthur M. Sackler Colloquia at the National Academy of Sciences pay tribute to this faith in communication as the prime mover of knowledge and culture.



Arthur M. Sackler Biography vii

Preface to the In the Light of Evolution Series xiii Preface to In the Light of Evoluion Volume II:

Biodiversity and Extinction xv



1 Ecological Extinction and Evolution in the Brave New Ocean 5 Jeremy B. C. Jackson

2 Are We in the Midst of the Sixth Mass Extinction?

A View from the World of Amphibians 27

David B. Wake and Vance T. Vredenburg

3 Patterns of Biodiversity and Endemism on Indo-West

Pacific Coral Reefs 45

Marjorie L. Reaka, Paula J. Rodgers, and Alexei U. Kudla

4 Homage to Linnaeus: How Many Parasites? How Many Hosts? 63 Andy Dobson, Kevin D. Lafferty, Armand M. Kuris,

Ryan F. Hechinger, and Walter Jetz ix x / Contents


5 Species Invasions and Extinction:

The Future of Native Biodiversity on Islands 85

Dov F. Sax and Steven D. Gaines

6 How Many Tree Species Are There in the Amazon and

How Many of Them Will Go Extinct? 107

Stephen P. Hubbell, Fangliang He, Richard Condit, Luis Borda-De-Âgua, James Kellner, and Hans ter Steege

7 Microbes on Mountainsides: Contrasting Elevational

Patterns of Bacterial and Plant Diversity 127

Jessica A. Bryant, Christine Lamanna, Hélène Morlon, Andrew J. Kerkhoff, Brian J. Enquist, and Jessica L. Green

8 Resistance, Resilience, and Redundancy in

Microbial Communities 149

Steven D. Allison and Jennifer B. H. Martiny


9 Extinction as the Loss of Evolutionary History 171 Douglas H. Erwin

10 Extinction and the Spatial Dynamics of Biodiversity 189 David Jablonski

11 Dynamics of Origination and Extinction in the

Marine Fossil Record 207

John Alroy

12 Megafauna Biomass Tradeoff as a Driver of

Quaternary and Future Extinctions 227

Anthony D. Barnosky

Contents / xi

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