Smithsonian Institution Scholarly Press smithsonian contributions to the marine sciences • number 39 Smithsonian Institution Scholarly Press Research and Discoveries The Revolution of Science through Scuba Edited by Michael A. Lang, Roberta L. Marinelli, Susan J. Roberts, and Phillip R. Taylor SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of “diffusing knowledge” was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: “It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge.” This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, com- mencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Botany Smithsonian Contributions to History and Technology Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Museum Conservation Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology In these series, the Institution publishes small papers and full-scale monographs that report on the research and collections of its various museums and bureaus. The Smithsonian Contributions Series are distributed via mailing lists to libraries, universities, and similar institu- tions throughout the world. Manuscripts submitted for series publication are received by the Smithsonian Institution Scholarly Press from authors with direct affilia- tion with the various Smithsonian museums or bureaus and are subject to peer review and review for compliance with manuscript preparation guidelines. General requirements for manuscript preparation are on the inside back cover of printed volumes. For detailed submissions require- ments and to review the “Manuscript Preparation and Style Guide for Authors,” visit the Submissions page at www.scholarlypress.si.edu. smithsonian contributions to the marine sciences • number 39 Research and Discoveries The Revolution of Science through Scuba Edited by Michael A. Lang, Roberta L. Marinelli, Susan J. Roberts, and Phillip R. Taylor WASHINGTON D.C. 2013 ABSTRACT Lang, Michael A., Roberta L. Marinelli, Susan J. Roberts, and Phillip R. Taylor, editors. Research and Discoveries: The Revo- lution of Science through Scuba. Smithsonian Contributions to the Marine Sciences, number 39, vi + 258 pages, 111 figures, 7 tables, 2013. The Smithsonian Institution, the National Science Foundation, and the Ocean Studies Board of the National Research Council convened the “Research and Discoveries: The Revolution of Science through Scuba” symposium on 24–25 May 2010 in Washington, D.C., to celebrate the advances and scientific contributions of research using self-contained un- derwater breathing apparatus (scuba). This volume presents 19 papers by 60 scholars of research findings, with particular focus on the scientific contributions accomplished using scuba. It is the first major effort to highlight and validate the use of scuba in science by evaluating the output of scientific research in high-impact journal publications. Thirteen papers report research findings and discoveries from around the world in environments such as coral reefs, oceanic blue water, under-ice polar habitats, and temperate kelp forests, providing perspectives on ecological scales and function, physiology, symbiosis and chemistry, biodiversity and behavior, and structured populations. The final six papers are illustrative of underwater research that was not only greatly facilitated by scuba, but could perhaps not have been accomplished without it. Topics range from biological studies on the coral holobiont to ecological roles of major algal groups on reefs and the functional role of small and cryptic metazoans. The research facilitated by scuba and reported in these papers focuses on the scientific results, not necessarily on the research methodologies using scuba to obtain those data and observations, and includes several case stud- ies. Where appropriate, laboratory studies complementary to underwater field observations are referenced. The symposium showed the strong integration and validation of scientific diving within the overall science domain since the introduction of scuba to the science community in 1951. Overarching symposium themes celebrated past, present, and future scientific diving contributions, and evaluated the accomplishments and impact of underwater research on the overall understanding of nature and its processes. Enduring materials from the symposium, including abstracts, speaker biographies, and webcast videos of presentations, have been posted at www.si.edu/sds. Cover images (left to right): Pseudopterogorgia elisabethae sea fan from Carrie Bow Cay, Belize (photo by Raphael Ritson- Williams); Macrocystis pyrifera kelp from San Diego, California (photo by Michael A. Lang); Leptonychotes weddelli seal from McMurdo Station, Antarctica (photo by Michael A. Lang). Published by SMITHSONIAN INSTITUTION SCHOLARLY PRESS P.O. Box 37012, MRC 957, Washington, D.C. 20013-7012 www.scholarlypress.si.edu Compilation copyright © 2013 Smithsonian Institution Text and images in this publication, including cover and interior designs, are owned either by the Smithsonian Institution, by contributing authors, or by third parties. Fair use of materials is permitted for personal, educational, or noncommercial purposes. Users must cite author and source of content, must not alter or modify copyrighted content, and must comply with all other terms or restrictions that may be applicable. Users are responsible for securing permission from a rights holder for any other use. Library of Congress Cataloging-in-Publication Data Research and discoveries : the revolution of science through scuba / edited by Michael A. Lang, Roberta L. Marinelli, Susan J. Roberts, and Phillip R. Taylor. pages cm. — (Smithsonian contributions to the marine sciences number ; 39) Includes bibliographical references. 1. Scuba diving. 2. Marine sciences. 3. Underwater exploration. I. Lang, Michael A. GV838.672.R47 2013 797.2’34—dc23 2013020367 ISSN: 0196-0768 (print); 1943-667X (online) The paper used in this publication meets the minimum requirements of the American National Standard for Per- manence of Paper for Printed Library Materials Z39.48–1992. Contents EXECUTIVE SUMMARY v THE REVOLUTION OF SCIENCE THROUGH SCUBA Scuba Revolutionizes Marine Science 3 Jon D. Witman, Paul K. Dayton, Suzanne N. Arnold, Robert S. Steneck, and Charles Birkeland Diving into the Past: Scuba and the Temporal Dimension of Coral Reefs 13 Ian G. Macintyre and Richard B. Aronson Diving Physiology and Decompression Sickness: Considerations from Humans and Marine Animals 23 Michael A. Lang, Jessica U. Meir, Tanya L. Streeter, and Karen B. Van Hoesen Saturation Diving and Underwater Laboratories: How Underwater Technology Has Aided Research on Coral Biology and Reef Ecology 39 Kenneth P. Sebens, Giacomo Bernardi, Mark R. Patterson, and Deron Burkepile Marine Chemical Ecology: A Science Born of Scuba 53 Joseph R. Pawlik, Charles D. Amsler, Raphael Ritson-Williams, James B. McClintock, Bill J. Baker, and Valerie J. Paul Scuba Diving in Blue Water: A Window on Ecology and Evolution in the Epipelagic Ocean 71 Laurence P. Madin, William M. Hamner, Steven H. D. Haddock, and George I. Matsumoto Scientific Polar Ice Diving 83 Adam G. Marsh, Amy L. Moran, and James B. McClintock The Use of Scuba to Study Early Life Histories of Marine Invertebrates 87 Harilaos A. Lessios, Howard R. Lasker, and Don R. Levitan Behavioral Ecology of Mobile Animals: Insights from In Situ Observations 99 David B. Eggleston, William F. Herrnkind, and Anson H. Hines Kelp Forests in California 115 Michael S. Foster, Daniel C. Reed, Mark H. Carr, Paul K. Dayton, Daniel P. Malone, John S. Pearse, and Laura Rogers-Bennett iv • SMITHSONIAN CONTRIBUTIONS to THE MARINE SCIENCES Kelpbeds as Classrooms: Perspectives and Lessons Learned 133 John S. Pearse, Mark H. Carr, Charles H. Baxter, James M. Watanabe, Michael S. Foster, Diana L. Steller, James A. Coyer, Brenda Konar, David O. Duggins, and Paul K. Dayton Rhodoliths and Rhodolith Beds 143 Michael S. Foster, Gilberto M. Amado Filho, Nicholas A. Kamenos, Rafael Riosmena-Rodríguez, and Diana L. Steller How Scuba Changed Our Understanding of Nature: Underwater Breakthroughs in Reef Fish Ecology 157 Mark H. Carr, Daniel P. Malone, Mark A. Hixon, Sally J. Holbrook, and Russell J. Schmitt RESEARCH FACILITATED BY SCUBA INTRODUCTION 171 Michael A. Lang, Roberta L. Marinelli, Susan J. Roberts, and Phillip R. Taylor Understanding the Coral Holobiont through Science and Scuba 173 Steve V. Vollmer, Andrew C. Baker, Mary-Alice Coffroth, C. Drew Harvell, and Mónica Medina The Nature of Macroalgae and Their Interactions on Reefs 187 Mark M. Littler and Diane S. Littler The Nature of Crustose Coralline Algae and Their Interactions on Reefs 199 Mark M. Littler and Diane S. Littler The Nature of Turf and Boring Algae and Their Interactions on Reefs 213 Mark M. Littler and Diane S. Littler The Ecological Significance of Sexual Reproduction by Tropical Green Algae 219 Kenneth E. Clifton Fine-Scale Interspecific Interactions on Coral Reefs: Functional Roles of Small and Cryptic Metazoans 229 Peter W. Glynn