WASHINGTON, D.C. 2009 Smithsonian at the Poles Contributions to International Polar Year Science Igor Krupnik, Michael A. Lang, and Scott E. Miller Editors A Smithsonian Contribution to Knowledge 00_FM_pg00i-xvi_Poles.indd i00_FM_pg00i-xvi_Poles.indd i 11/17/08 8:41:31 AM11/17/08 8:41:31 AM This proceedings volume of the Smithsonian at the Poles symposium, sponsored by and convened at the Smithsonian Institution on 3?4 May 2007, is published as part of the International Polar Year 2007?2008, which is sponsored by the International Council for Science (ICSU) and the World Meteorological Organization (WMO). Published by Smithsonian Institution Scholarly Press P.O. Box 37012 MRC 957 Washington, D.C. 20013-7012 www.scholarlypress.si.edu Text and images in this publication may be protected by copyright and other restrictions or owned by individuals and entities other than, and in addition to, the Smithsonian Institution. Fair use of copyrighted material includes the use of protected materials for personal, educational, or noncommercial purposes. Users must cite author and source of content, must not alter or modify content, and must comply with all other terms or restrictions that may be applicable. Cover design: Piper F. Wallis Cover images: (top left) Wave-sculpted iceberg in Svalbard, Norway (Photo by Laurie M. Penland); (top right) Smithsonian Scientifi c Diving Offi cer Michael A. Lang prepares to exit from ice dive (Photo by Adam G. Marsh); (main) Kongsfjorden, Svalbard, Norway (Photo by Laurie M. Penland). Library of Congress Cataloging-in-Publication Data Smithsonian at the poles : contributions to International Polar Year science / Igor Krupnik, Michael A. Lang, and Scott E. Miller, editors. p. cm. ISBN 978-0-9788460-1-5 (pbk. : alk. paper) 1. International Polar Year, 2007?2008. 2. Polar regions?Research?Congresses. 3. Research?Polar regions?Congresses. 4. Arctic regions?Research?Congresses. 5. Antarctica?Research?Congresses. 6. Polar regions?Environmental conditions?Congresses. 7. Climatic changes?Detection?Polar regions?Congresses. I. Krupnik, Igor. II. Lang, Michael A. III. Miller, Scott E. G587.S65 2009 559.8?dc22 2008042055 ISBN-13: 978-0-9788460-1-5 ISBN-10: 0-9788460-1-X The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48?1992. 00_FM_pg00i-xvi_Poles.indd ii00_FM_pg00i-xvi_Poles.indd ii 11/17/08 8:41:32 AM11/17/08 8:41:32 AM FOREWORD by Ira Rubinoff ix EXECUTIVE SUMMARY by Michael A. Lang xii INTRODUCTION by Igor Krupnik, Michael A. Lang, and Scott E. Miller xiv IPY HISTORIES AND LEGACIES Advancing Polar Research and Communicating Its Wonders: Quests, Questions, and Capabilities of Weather and Climate Studies in International Polar Years 1 James R. Fleming, Colby College Cara Seitchek, Woodrow Wilson International Center for Scholars Cooperation at the Poles? Placing the First International Polar Year in the Context of Nineteenth-Century Scientifi c Exploration and Collaboration 13 Marc Rothenberg, National Science Foundation The Policy Process and the International Geophysical Year, 1957?1958 23 Fae L. Korsmo, National Science Foundation Preserving the Origins of the Space Age: The Material Legacy of the International Geophysical Year (1957?1958) at the National Air and Space Museum 35 David H. DeVorkin, National Air and Space Museum, Smithsonian Institution From Ballooning in the Arctic to 10,000-Foot Runways in Antarctica: Lessons from Historic Archaeology 49 Noel D. Broadbent, National Museum of Natural History, Smithsonian Institution Contents 00_FM_pg00i-xvi_Poles.indd iii00_FM_pg00i-xvi_Poles.indd iii 11/17/08 8:41:33 AM11/17/08 8:41:33 AM CULTURAL STUDIES ?Of No Ordinary Importance?: Reversing Polarities in Smithsonian Arctic Studies 61 William W. Fitzhugh, National Museum of Natural History, Smithsonian Institution Yup?ik Eskimo Contributions to Arctic Research at the Smithsonian 79 Ann Fienup-Riordan, National Museum of Natural History Arctic Studies Center, Anchorage Smithsonian Contributions to Alaskan Ethnography: The First IPY Expedition to Barrow, 1881?1883 89 Ernest S. Burch Jr., National Museum of Natural History Arctic Studies Center, Camp Hill The Art of I?upiaq Whaling: Elders? Interpretations of International Polar Year Ethnological Collections 99 Aron L. Crowell, National Museum of Natural History Arctic Studies Center, Anchorage From Tent to Trading Post and Back Again: Smithsonian Anthropology in Nunavut, Nunavik, Nitassinan, and Nunatsiavut?The Changing IPY Agenda, 1882?2007 115 Stephen Loring, National Museum of Natural History, Smithsonian Institution ?The Way We See It Coming?: Building the Legacy of Indigenous Observations in IPY 2007?2008 129 Igor Krupnik, National Museum of Natural History, Smithsonian Institution SYSTEMATICS AND BIOLOGY OF POLAR ORGANISMS Species Diversity and Distributions of Pelagic Calanoid Copepods from the Southern Ocean 143 E. Taisoo Park, Texas A&M University Frank D. Ferrari, National Museum of Natural History, Smithsonian Institution Brooding and Species Diversity in the Southern Ocean: Selection for Brooders or Speciation within Brooding Clades? 181 John S. Pearse, University of California, Santa Cruz Richard Mooi, California Academy of Sciences Susanne J. Lockhart, California Academy of Sciences Angelika Brandt, Zoologisches Institut und Zoologisches Museum, Hamburg Persistent Elevated Abundance of Octopods in an Overfi shed Antarctic Area 197 Michael Vecchione, National Marine Fisheries Service Louise Allcock, Queen?s University Belfast Uwe Piatkowski, Universitat Kiel, Germany iv SMITHSONIAN AT THE POLES 00_FM_pg00i-xvi_Poles.indd iv00_FM_pg00i-xvi_Poles.indd iv 2/4/09 1:00:43 PM2/4/09 1:00:43 PM Elaina Jorgensen, Alaska Fisheries Science Center Iain Barratt, Queen?s University Belfast Cold Comfort: Systematics and Biology of Antarctic Bryozoans 205 Judith E. Winston, Virginia Museum of Natural History Considerations of Anatomy, Morphology, Evolution, and Function for Narwhal Dentition 223 Martin T. Nweeia, Harvard School of Dental Medicine Cornelius Nutarak, Elder, Community of Mittimatilik, Nunavut Frederick C. Eichmiller, Delta Dental of Wisconsin Naomi Eidelman, ADAF Paffenbarger Research Center Anthony A. Giuseppetti, ADAF Paffenbarger Research Center Janet Quinn, ADAF Paffenbarger Research Center James G. Mead, National Museum of Natural History, Smithsonian Institution Kaviqanguak K?issuk, Hunter, Community of Qaanaaq, Greenland Peter V. Hauschka, National Museum of Natural History, Smithsonian Institution Ethan M. Tyler, National Institutes of Health Charles Potter, National Museum of Natural History, Smithsonian Institution Jack R. Orr, Fisheries and Oceans, Canada, Arctic Research Division Rasmus Avike, Hunter, Community of Qaanaaq, Greenland Pavia Nielsen, Elder, Community of Uummannaq, Greenland David Angnatsiak, Elder, Community of Mittimatilik, Nunavut METHODS AND TECHNIQUES OF UNDER-ICE RESEARCH Scientifi c Diving Under Ice: A 40-Year Bipolar Research Tool 241 Michael A. Lang, Offi ce of the Under Secretary for Science, Smithsonian Institution Rob Robbins, Raytheon Polar Services Company Environmental and Molecular Mechanisms of Cold Adaptation in Polar Marine Invertebrates 253 Adam G. Marsh, University of Delaware, Lewes Milestones in the Study of Diving Physiology: Antarctic Emperor Penguins and Weddell Seals 265 Gerald Kooyman, Scripps Institution of Oceanography Interannual and Spatial Variability in Light Attenuation: Evidence from Three Decades of Growth in the Arctic Kelp, Laminaria solidungula 271 Kenneth H. Dunton, University of Texas Marine Science Institute Susan V. Schonberg, University of Texas Marine Science Institute Dale W. Funk, LGL Alaska Research Associates, Inc. Life under Antarctic Pack Ice: A Krill Perspective 285 Langdon B. Quetin, University of California, Santa Barbara Robin M. Ross, University of California, Santa Barbara CONTENTS v 00_FM_pg00i-xvi_Poles.indd v00_FM_pg00i-xvi_Poles.indd v 11/17/08 8:41:34 AM11/17/08 8:41:34 AM ENVIRONMENTAL CHANGE AND POLAR MARINE ECOSYSTEMS Inhibition of Phytoplankton and Bacterial Productivity by Solar Radiation in the Ross Sea Polynya 299 Patrick J. Neale, Smithsonian Environmental Research Center Wade H. Jeffrey, University of West Florida Cristina Sobrino, University of Vigo, Spain J. Dean Pakulski, University of West Florida Jesse Phillips-Kress, Smithsonian Environmental Research Center Amy J. Baldwin, Florida Department of Environmental Protection Linda A. Franklin, Smithsonian Environmental Research Center Hae-Cheol Kim, Harte Research Institute Southern Ocean Primary Productivity: Variability and a View to the Future 309 Walker O. Smith Jr., Virginia Institute Marine Sciences Josefi no C. Comiso, NASA Goddard Space Flight Center Chromophoric Dissolved Organic Matter Cycling during a Ross Sea Phaeocystis antarctica Bloom 319 David J. Kieber, SUNY College of Environmental Science and Forestry Dierdre A. Toole, Woods Hole Oceanographic Institution Ronald P. Kiene, University of South Alabama Capital Expenditure and Income (Foraging) during Pinniped Lactation: The Example of the Weddell Seal (Leptonychotes weddellii) 335 Regina Eisert, National Zoological Park, Smithsonian Institution Olav T. Oftedal, Smithsonian Environmental Research Center Latitudinal Patterns of Biological Invasions in Marine Ecosystems: A Polar Perspective 347 Gregory M. Ruiz, Smithsonian Environmental Research Center Chad L. Hewitt, Australian Maritime College POLAR ASTRONOMY: OBSERVATIONAL COSMOLOGY Cosmology from Antarctica 359 Robert W. Wilson, Smithsonian Astrophysical Observatory Antony A. Stark, Smithsonian Astrophysical Observatory Feeding the Black Hole at the Center of the Milky Way: AST/RO Observations 370 Christopher L. Martin, Oberlin College HEAT: The High Elevation Antarctic Terahertz Telescope 373 Christopher K. Walker, University of Arizona Craig A. Kulesa, University of Arizona vi SMITHSONIAN AT THE POLES 00_FM_pg00i-xvi_Poles.indd vi00_FM_pg00i-xvi_Poles.indd vi 11/17/08 8:41:34 AM11/17/08 8:41:34 AM Watching Star Birth from the Antarctic Plateau 381 Nick F. H. Tothill, University of Exeter Mark J. McCaughrean, University of Exeter Christopher K. Walker, University of Arizona, Tucson Craig Kulesa, University of Arizona, Tucson Andrea Loehr, Smithsonian Astrophysical Observatory Stephen Parshley, Cornell University Antarctic Meteorites: Exploring the Solar System from the Ice 387 Timothy J. McCoy, National Museum of Natural History, Smithsonian Institution Linda C. Welzenbach, National Museum of Natural History, Smithsonian Institution Catherine M. Corrigan, National Museum of Natural History, Smithsonian Institution INDEX 395 CONTENTS vii 00_FM_pg00i-xvi_Poles.indd vii00_FM_pg00i-xvi_Poles.indd vii 11/17/08 8:41:35 AM11/17/08 8:41:35 AM 00_FM_pg00i-xvi_Poles.indd viii00_FM_pg00i-xvi_Poles.indd viii 11/17/08 8:41:35 AM11/17/08 8:41:35 AM Foreword O n behalf of Smithsonian colleagues and as a tropical biologist, I ex- tend a warm welcome to this International Polar Year 2007?2008 science symposium. The commonality between poles and tropics is their shared image of ?remoteness? that keeps them removed from the thoughts of society that sponsors our research. In fact, we believe that the poles and tropics are now ?canaries in the coalmine,? because they are at the forefront of global change. We face a desperate task of educating society that our global problems are not restricted to the densely populated areas of the United States, Europe, and Asia. We must increase our understanding of how polar regions affect the habit- ability of our planet through long-term monitoring and observations of short- term cyclical changes, geosphere/atmosphere interactions, and interconnectivity of physical, biological, and social systems. Coming to Washington in 2007 to adopt a broader set of science responsi- bilities than I previously had at the Smithsonian Tropical Research Institute, I have found a multitude of polar interests throughout the Smithsonian. I high- light the following Smithsonian programs, which you will learn more about during this Smithsonian symposium. The U.S. Antarctic Meteorite Program is headed by Tim McCoy, Depart- ment of Mineral Sciences, National Museum of Natural History. Since 1976, Smithsonian, NSF and NASA have supported the accession of over 12,000 me- teorite specimens from the Antarctic ice sheets in an attempt to better under- stand the history of our solar system. The U.S. Antarctic Program Invertebrate Collections are deposited in the National Museum of Natural History and currently number over 900,000 speci- mens. Archival samples from the Palmer Long-Term Ecological Research site are now also included. Rafael Lemaitre, Chairman of the Invertebrate Zoology De- partment, reports that this program (co-sponsored by SI and NSF) has loaned over 170,000 specimens in 138 separate lots to researchers in 22 countries since 1995. Antarctic photobiology has been co-sponsored by NSF and the Smithsonian Environmental Research Center since 1990. Patrick Neale, Principal Investiga- tor, investigates the colonial alga (Phaeocystis antarctica) that dominates spring 00_FM_pg00i-xvi_Poles.indd ix00_FM_pg00i-xvi_Poles.indd ix 11/17/08 8:41:35 AM11/17/08 8:41:35 AM blooms in a polynya well within the ozone hole, expos- ing plankton to elevated UV-B. The continuous daylight characteristic of this time of year has implications for the regulation of DNA repair, most of which normally occurs at night. The Northern Latitudes Invasions Biology Program, directed by Gregory Ruiz of the Smithsonian Environ- mental Research Center has documented the northward spread into six Alaskan regions of several nonnative spe- cies. Emerging data suggest that polar systems are cer- tainly vulnerable to invasions. Experimental analyses and modeling of the environmental tolerance of known non- native species and their capacity to colonize polar systems are underway. The National Zoological Park?s Olav Oftedal is study- ing Weddell Seal energetics in Antarctica. This study of seal capital expenditure (reliance on stored reserves), lac- tation energetics and the importance of food intake relies on a novel multimarker approach. The relative importance of these expenditures and energy transfer to pups, in the evolution of a mixed capital and income breeding strategy, is being evaluated. The NSF?s U.S. Antarctic Diving Program has been managed since 2001 by Michael Lang in the Smithsonian?s Offi ce of the Under Secretary for Science through an Inter- agency Agreement. During this period the program reports an average of 35 scientifi c divers per year logging a total of over 4,800 under-ice dives while enjoying a remarkable safety record and scientifi c productivity. Smithsonian ice diving courses are taught regularly at the Svalbard Arctic Marine Laboratory. Astrophysical results include fi rst light achieved with the NSF 10-m South Pole Telescope, 16 February 2007, obtaining maps of Jupiter at wavelengths at 2 mm and 3 mm. The Antarctic Submillimeter Telescope and Remote Observatory was operated by Antony Stark of the Smith- sonian Astrophysical Observatory since 1995 as part of the Center for Astrophysical Research in Antarctica, un- der NSF agreement. The Arctic Studies Center of the National Museum of Natural History was established by the Department of Anthropology?s William Fitzhugh in 1988, with a second offi ce operating since 1995 in Anchorage. Its focus is on cultural heritage studies and indigenous knowledge of sea ice, marine mammals and Arctic climate change. The re- cent exhibit ?Arctic: A Friend Acting Strangely? premiered in 2006 at the Natural History Museum. The United States marked the start of International Polar Year (IPY) 2007?2008 with an opening event hosted by the National Academies and the National Sci- ence Foundation on 26 February 2007. IPY is a global research effort to better understand the polar regions and their climatic effect on Earth. The research completed during IPY 2007?2008 will provide a baseline for under- standing future environmental change. Smithsonian at the Poles: Contributions to International Polar Year Science is one of the major inaugural science symposia of this IPY. It is also one of the fi rst efforts in disseminating scientifi c knowledge and research inspired by IPY to be undertaken and published during this IPY period. Another Smithsonian-initiated conference, Making Science Global: Reconsidering the Social and Intellectual Implications of the International Polar and Geophysical Years, will convene in November 2007. Making Science Global, an NSF-supported conference, explores the impe- tus for (and the impact upon) science, society, and culture of the IPYs of 1882?1883 and 1932?1933, and the In- ternational Geophysical Year of 1957?1958. It is devoted to sharing historical perspectives that might be useful to those involved in the current IPY. Sessions will explore the origins of these campaigns, their political dimensions, and their consequences. Specifi c themes include the place of the poles in human imagination, the role of polar exploration in discipline formation, cultural nationalism, politics, and transnationality. Additionally, there are sessions planned on the emergence of the modern geosciences, the uses of new technologies to explore the poles, changing assess- ments of the nature of human cultures in high latitudes, and polar contributions to environmental awareness. The fi nal session of the conference, Polar History: Perspectives on Globalization in the Geosciences, is a plenary session at the annual meeting of the History of Science Society. The Antarctic Treaty Summit: Science-Policy Interac- tions in International Governance will be co-sponsored by the Smithsonian Institution and will be convened at the National Museum of Natural History from 30 Novem- ber through 3 December 2009. This summit celebrates the fi ftieth anniversary of the signature-day for the Antarctic Treaty in the city where it was adopted ?in the interest of all mankind.? I would like to thank the NSF Offi ce of Polar Pro- grams for its support of this symposium and the Sympo- sium Committee Michael Lang, Scott Miller, Igor Krupnik, Bill Fitzhugh, Rafael Lemaitre, Pat Neale, and Tony Stark and the symposium speakers for their efforts. Ira Rubinoff Smithsonian Institution Acting Under Secretary for Science 3 May 2007 x SMITHSONIAN AT THE POLES 00_FM_pg00i-xvi_Poles.indd x00_FM_pg00i-xvi_Poles.indd x 11/17/08 8:41:35 AM11/17/08 8:41:35 AM Executive Summary S mithsonian at the Poles: Contributions to International Polar Year Sci- ence presents the proceedings of an interdisciplinary symposium dedi- cated to the opening of International Polar Year (IPY) 2007?2008 and hosted by the Smithsonian Institution on 3?4 May 2007. The volume refl ects partnerships across various Smithsonian research units engaged in polar research as well as collaboration with U.S. government agencies with active IPY 2007?2008 programs, including the National Science Foundation (NSF), the National Aeronautics and Space Administration (NASA), the National Oceanic and Atmospheric Administration (NOAA), and the Department of the Interior (DOI). Smithsonian at the Poles is the fi rst of many publications by Smithsonian scientists and curators and their collaborators envisioned in association with U.S. scholarly and public programs for IPY. 1 The Smithsonian at the Poles symposium was convened by the Smithsonian Institution?s Offi ce of the Under Secretary for Science (OUSS) with major sup- port from NSF. The three symposium co-Chairs?Igor Krupnik, Department of Anthropology, National Museum of Natural History (NMNH), and Michael Lang and Scott Miller, both of OUSS?relied on support from many scientists and curators active in polar research in both the Arctic and Antarctic regions. The Symposium Committee included the three co-Chairs as well as Antony Stark (Smithsonian Astrophysical Observatory), William Fitzhugh (Department of Anthropology, NMNH), Rafael Lemaitre (Department of Invertebrate Zool- ogy, NMNH), and Patrick Neale (Smithsonian Environmental Research Center). The committee members were designated Chairs of the individual symposium sessions and coordinators of the corresponding sections in this volume. On May 3, 2007, Paul Risser, then Acting Director of NMNH, opened the Smithsonian at the Poles symposium. The fi rst plenary session was concluded with an address by Ira Rubinoff, then Acting Under Secretary for Science. The agenda included plenary session presentations by Robert W. Corell (Heinz Cen- ter), Robert W. Wilson (SAO), Donal T. Manahan (University of Southern Cali- fornia) and William W. Fitzhugh (NMNH); six concurrent thematic sessions (IPY Histories and Legacies, People and Cultures, Systematics and Biology of Polar Organisms, Methods and Techniques of Under-Ice Research, Environmental 00_FM_pg00i-xvi_Poles.indd xi00_FM_pg00i-xvi_Poles.indd xi 11/17/08 8:41:36 AM11/17/08 8:41:36 AM Change and Polar Marine Ecosystems, and Polar Astron- omy: Observational Cosmology) and panel discussions; an evening public cinematic event presented by Adam Ravetch and Norbert Wu; and a keynote presentation by James W. C. White (University of Colorado, Boulder). The fi rst ever Smithsonian polar science assembly that covered both the Arctic and Antarctica, Smithsonian at the Poles featured more than 200 scholars, members of the public, agency rep- resentatives from the Smithsonian, NSF, NOAA, numerous U.S. universities, Australia, and Germany. The symposium was originally pledged in 2005 by the Offi ce of the Under Secretary for Science in addition to an educational exhibit on Arctic climate change, Arctic: A Friend Acting Strangely (2006), access to the Smithson- ian polar collections, and the use of the Smithsonian fa- cilities for IPY-related public activities (Anonymous, 2005; Krupnik, 2006). The Smithsonian Institution is pleased to contribute this summary of the Institution?s research around the Poles. 2 ACKNOWLEDGMENTS The National Science Foundation contributed funding in support of this polar science symposium under Grant No. 0731478 to Michael A. Lang, Principal Investigator, Smithsonian Offi ce of the Under Secretary for Science. NOTES 1. The Smithsonian at the Poles symposium was part of the IPY 2007?2008, a joint initiative of the International Council for Sciences (ICSU) and the World Meteorological Organization (WMO). Scholars from more than 60 nations participated in some 250 projects under the IPY 2007?2008 science program. 2. More details related to the symposium program are available on the symposium website at www.si.edu/ipy, the offi cial U.S. site www.ipy. gov, in Smithsonian Institution press releases, and in the news media. LITERATURE CITED Anonymous. 2005. U.S. Arctic Research Plan. Biennial Revision: 2006? 2010. Arctic Research of the United States, 19 (Fall?Winter):20? 21. Krupnik, I. 2006. Smithsonian Institution. Arctic Research of the United States, 20:142?146. xii SMITHSONIAN AT THE POLES 00_FM_pg00i-xvi_Poles.indd xii00_FM_pg00i-xvi_Poles.indd xii 11/17/08 8:41:36 AM11/17/08 8:41:36 AM ?Smithsonian at the Poles?: A 150-Year Venture T he Smithsonian Institution has a strong legacy of International Polar Year (IPY) activities dating to the fi rst IPY in 1882?1883. The fi rst two Smithsonian Secretaries, Joseph Henry (1846?1878) and Spencer F. Baird (1878?1887), were strong proponents of the advancement of science in the polar regions and of the disciplines that eventually formed the core of the program for that fi rst IPY: meteorology, astronomy, geology, and natural history, as well as studies of the polar residents and their cultures. By the fi rst IPY, the Smithsonian had forged partnerships with federal agen- cies, private organizations, and individual explorers active in polar regions (Baird, 1885a; 1885b). For instance, the Signal Offi ce of the then U.S. War Department was in charge of preparations for U.S. IPY missions to Barrow (1881?1883) and Lady Franklin Bay (1881?1884), supported by expedition scientists whom the Smithsonian helped to select and train. The Institution also offered its facilities and libraries, and the expertise of its curators to the returning IPY parties and was granted most of the American IPY-1 natural science and ethnological col- lections, expedition photographs, and personal memorabilia returning from the North. As a result, the Smithsonian?s early natural history collections from Bar- row, Alaska, are among the most comprehensive in the world, as are those from Labrador, accessioned in 1884. The Smithsonian published two monographs as contributions to its Annual Reports of the Bureau of Ethnology series and many shorter papers that described the collections accessioned from the IPY missions to Alaska and Canada (Murdoch, 1892; Turner, 1894; Fitzhugh, 1988; Loring, 2001). These Smithsonian holdings are now a source of knowledge to scientists and cultural information to indigenous communities, who view the early IPY collections as their prime heritage resource. The Smithsonian?s involvement in the second International Polar Year 1932? 1933 and in the International Geophysical Year (IGY) 1957?1958 (originally planned as the ?third IPY?) was modest, although the National Air and Space 00_FM_pg00i-xvi_Poles.indd xiii00_FM_pg00i-xvi_Poles.indd xiii 11/17/08 8:41:36 AM11/17/08 8:41:36 AM Museum (NASM) houses a substantial collection related to the space explorations from the IGY era. Since the 1970s, the Smithsonian has been actively engaged in Antarctic research, initially through hosting (since 1976) the U.S. Antarctic Meteorite Program, a co- operative effort with NASA and NSF aimed at collection, curation, and long-term storage of meteorites recovered from the Antarctic ice sheets by U.S. scientists. Today, curators of the NMNH Department of Mineral Sciences classify each of the meteorites accessioned and publish the results in the Antarctic Meteorite Newsletter, issued bian- nually by NASA?s Johnson Space Center. For decades, the Smithsonian has focused socio- cultural and heritage studies on the indigenous people of the Arctic, supported by its long-established tradition of cultural research in the North and its northern ethno- graphic collections from Alaska, Canada, Greenland, and Siberia. Since 1988, these efforts have been spearheaded through the creation of the Smithsonian?s polar cultural studies unit, the Arctic Studies Center (ASC) at NMNH?s Department of Anthropology. Under a cooperative agree- ment with the Anchorage Museum of History and Art, the ASC operates its Alaskan regional offi ce in Anchorage. In the 1990s, NMNH agreed to host the national Antarctic invertebrate collection for the NSF United States Antarctic Program (USAP), which now also incorpo- rates the Palmer Long-Term Ecological research (LTER) voucher specimens. Now totaling over 900,000 specimens, 170,000 specimens have been loaned in 138 separate lots to polar researchers in 22 countries. The precursor of this collection was the Smithsonian Oceanographic Sorting Center, which processed over 38 million polar specimens and distributed them to the scientifi c community between 1965 and 1992 (Moser and Nicol, 1997). One of the Smithsonian?s long-term polar ventures is its astrophysical projects at the South Pole station in Antarctica. The Smithsonian Astrophysical Observatory?s (SAO) Antarctic projects have included the Antarctic Sub- millimeter Telescope and Remote Observatory (AST/RO) operated by SAO 1995?2007 as part of the Center for Astrophysical Research in Antarctica, under NSF agree- ment. Now SAO collaborates on several projects using the newly built South Pole Telescope (SPT), a 10-m diameter telescope for millimeter and submillimeter observations. The SPT holds the promise of making a signifi cant break- through in our understanding of the universe and of phys- ics in general by surveying the entire southern sky, one- third of the celestial sphere, and potentially discovering 30,000 new clusters of galaxies during the next two to three years. This data will provide substantially improved measures of ?dark energy,? the newly discovered force that is driving the acceleration of the universe. Since 2001, the Smithsonian Scientifi c Diving Program (SDP), established by the Offi ce of the Under Secretary for Science in 1990, has managed the NSF Offi ce of Polar Pro- grams?sponsored scientifi c diving activities at the U.S. Antarctic McMurdo and Palmer Stations and from the re- search vessels L.M. Gould and N.B. Palmer. On average, 35 scientists have dived under ice each year through USAP dive program. More than 4,800 scientifi c ice dives were logged during 2000?2005. Formal ice diving training is provided by the SDP through biannual ice diving courses in Ny-?lesund, Svalbard. The USAP scientifi c diving ex- posures enjoy a remarkable safety record and proven sci- entifi c productivity as an underwater research tool. Polar diving history spans only 60 years, since the United States? fi rst major post-war Antarctic venture and the invention of the scuba regulator, and was thus not represented well until this fourth IPY. Smithsonian scientists are actively engaged in IPY 2007?2008 projects from astrophysical observations at the South Pole to the use of Smithsonian collections in educational and knowledge preservation programs in in- digenous communities across the Arctic. Ongoing projects include: the Photobiology and Solar Radiation Antarctic Research Program, supported by the Smithsonian Envi- ronmental Research Center (SERC) and NSF; Northern Latitudes Invasions Biology (NLIB) by SERC?s Invasions Biology Program; NMNH?s professional collections man- agement services provided for the NSF USAP and the inter- national scientifi c community through the ?USNM Polar Invertebrate Online Databases?; investigations of Weddell seal energetics, supported by NSF and conducted by the Smithsonian?s National Zoological Park; and bipolar in- ternational polar diving safety research in Svalbard and McMurdo Station (Lang and Sayer, 2007). For the fourth IPY in 2007?2008, there was Smithso- nian representation in all interagency planning meetings. Igor Krupnik served on the fi rst U.S. National IPY Com- mittee in 2003?2005 (NAS, 2004) and since 2004 on the Joint Committee for IPY 2007?2008, the international steering body that supervises planning and implementa- tion, to represent social and human studies. By its very nature, each International Polar Year is an invitation to the history of science, polar research, and the legacy of polar exploration. Launched approximately ev- ery 50 years (or after 25 years in the case of International Geophysical Year 1957?1958), these international ventures create incentives for scientists to test earlier records and to revisit the studies of their predecessors. Each new initiative xiv SMITHSONIAN AT THE POLES 00_FM_pg00i-xvi_Poles.indd xiv00_FM_pg00i-xvi_Poles.indd xiv 11/17/08 8:41:37 AM11/17/08 8:41:37 AM offers an unparalleled vantage point into the advancement of science, scholarly planning, and collaboration since the previous IPY. The ?Smithsonian at the Poles? symposium of 2007 and this resulting volume are the most recent Smithsonian contributions to the ever-growing infl uence and inspiration of the International Polar Year. LITERATURE CITED Baird, S. F. 1885a. Report of Professor Baird, Secretary of the Smithson- ian Institution, for 1883. Annual Report of the Board of Regents of the Smithsonian Institution, Showing the Operations, Expendi- tures, and Condition of the Institution for the Year 1883. Washing- ton, D.C.: Government Printing Offi ce. ???. 1885b. Report of Professor Baird, Secretary of the Smithsonian Institution, for 1884. Annual Report of the Board of Regents of the Smithsonian Institution, Showing the Operations, Expenditures, and Condition of the Institution for the Year 1884. Washington, D.C.: Government Printing Offi ce. Fitzhugh, W. W. 1988. ?Introduction to the 1988 Edition.? In Ethnologi- cal Results of the Point Barrow Expedition, ed. John Murdoch, pp. xiii?xlix. Washington, D.C.: Smithsonian Institution Press. Lang, M. A., and M. D. J. Sayer, eds. 2007. Proceedings of the Inter- national Polar Diving Workshop. Svalbard, 15?21 March 2007. Washington, D.C.: Smithsonian Institution. Loring, S. 2001. Introduction to Lucien M. Turner and the Beginnings of Smithsonian Anthropology in the North. In Ethnology of the Ungava District, Hudson Bay Territory, L. M. Turner, pp. vii?xxxii. Washington, D.C.: Smithsonian Institution Press. Moser, W. E., and J. Nicol. 1997. The National Museum of Natural His- tory. Antarctic Journal of the United States, 23:11?16. Murdoch, J. 1892. ?Ethnological Results of the Point Barrow Expedi- tion.? In Ninth Annual Report of the Bureau of Ethnology 1887? 88, pp. 1?441. Washington, D.C.: Government Printing Offi ce (2nd ed., 1988, Smithsonian Institution Press). National Academy of Sciences Polar Research Board. 2004. A Vision for the International Polar Year 2007?2008. Washington, D.C.: Na- tional Academies Press. Turner, L. 1894. ?Ethnology of the Ungava District, Hudson Bay Ter- ritory.? In Eleventh Annual Report of the Bureau of Ethnology, 1889?1890, ed. J. W. Powell, pp.159?350. Washington, D.C.: Gov- ernment Printing Offi ce. INTRODUCTION xv 00_FM_pg00i-xvi_Poles.indd xv00_FM_pg00i-xvi_Poles.indd xv 11/17/08 8:41:37 AM11/17/08 8:41:37 AM 00_FM_pg00i-xvi_Poles.indd xvi00_FM_pg00i-xvi_Poles.indd xvi 11/17/08 8:41:37 AM11/17/08 8:41:37 AM ABSTRACT. Since its inception, the Smithsonian Institution has been a leader in ad- vancing science and communicating its wonders. It functioned as a ?national center for atmospheric research? in the nineteenth century and served as a model for the founding of the U.S. Weather Bureau. Its archives and collections document Smithsonian support and involvement over the years in many of the early weather and climate science ini- tiatives: in both the fi rst and second International Polar Years; in the founding of the Arctic Institute of North America and the National Academy of Sciences Conference on the Antarctic; and in the International Geophysical Year in 1957? 1958. This presenta- tion examines science, technology, and public opinion surrounding weather and climate research at both poles, from the middle of the nineteenth century through the fi rst and second International Polar Years and the International Geophysical Year, up to the cur- rent International Polar Year 2007? 2008. INTRODUCTION During the past two centuries, the scientifi c study of weather and climate has changed repeatedly and dramatically. In different eras, telegraphy, radio, rocketry, electronic computing, and satellite meteorology have provided new capabilities for measuring, monitoring, modeling, and theorizing about the atmosphere. While the scale and sophistication has changed, what has not changed is the need for coop- erative efforts spanning the largest areas possible? including the poles. Over the years, polar science has served as a very positive example of international peaceful cooperation. The fi rst International Polar Year (IPY-1) of 1882? 1883 involved 11 nations in a coordinated effort to study atmospheric changes and ?electrical weather? as shown by magnetic disturbances and the polar lights. These efforts were confi ned to surface observations (Heathcote and Armitage, 1959). In IPY-2 of 1932? 1933, 40 nations were involved in a global program to study meteorol- ogy, magnetism, and radio science as related to the ionosphere, using instrumented balloons to reach altitudes as high as 10 kilometers (Laursen, 1959). The Inter- national Geophysical Year (IGY) of 1957? 1958 involved 67 nations in what the British astronomer and geophysicist Sydney Chapman (1888? 1970) called, ?the common study of our planet by all nations for the benefi t of all.? Using rockets, new earth-orbiting satellites, and a variety of other techniques, scientists studied James R. Fleming, Science, Technology, and Society Program, Colby College, 5881 May- fl ower Hill, Waterville, ME 04901, USA. Cara Seitchek, Woodrow Wilson International Center for Scholars, 1300 Pennsylvania Avenue, NW, Washington, DC 20004-3027, USA. Correspond- ing author: J. R. Fleming (jfl eming@colby.edu). Accepted 29 May 2008. Advancing Polar Research and Communicating Its Wonders: Quests, Questions, and Capabilities of Weather and Climate Studies in International Polar Years James R. Fleming and Cara Seitchek 01_Fleming_pg001-012_Poles.indd 101_Fleming_pg001-012_Poles.indd 1 11/17/08 8:31:07 AM11/17/08 8:31:07 AM 2 SMITHSONIAN AT THE POLES / FLEMING AND SEITCHEK the interaction of the sun and the earth, with a special focus on Antarctica (Chapman, 1959a:102). These international cooperative efforts serve as benchmarks for meteorological efforts in high latitudes and help reveal larger issues con- cerning the continuity and interconnectedness of the science and technology of weather and climate research. Indeed, each successive IPY was based upon the technological in- novations of its era and was informed by cutting-edge scien- tifi c theories and hypotheses. The launch of the current IPY of 2007? 2008, involving more than 60 nations, provides an occasion to look back and to look beyond for larger mes- sages about weather and climate research, the interrelation- ships created by international science, and the connection between science, technology, and popular culture. HISTORICAL PRECEDENTS Cooperative scientifi c observations date to the early seventeenth century. In the closing decades of the eigh- teenth century in Europe, and slightly later in Russia and the United States, serious attempts were made to broaden the geographic coverage of weather observations, stan- dardize their collection, and publish the results. Individual observers in particular locales dutifully tended to their journals, and networks of cooperative observers gradually extended the meteorological frontiers. A century before IPY-1, the Societas Meteorologica Palatina (1781? 1795), an international organization whose members represented the chief European scientifi c institutions, collected obser- vations from a network of 57 stations extending from Si- beria to North America and southward to the Mediterra- nean. The observers, who received instruments, forms, and instructions free of charge, sent their results to Mannheim, Germany, where they were published in extenso (Cassidy, 1985:8? 25; Societas Meteorologica Palatina, 1783? 1795). Many subsequent projects emulated their example. In the 1830s Sir John Herschel (1791? 1872), then in Cape Town, South Africa, initiated the practice of col- lecting extensive hourly geophysical measurements on ?term days?? 36-hour periods surrounding the dates of the equinoxes and solstices. The measurements, accord- ing to a common plan, were taken simultaneously from widely dispersed stations in order to obtain knowledge of the ?correspondence of [the] movements and affec- tions [of the atmosphere] over great regions of the earth?s surface, or even over the whole globe? (Herschel, 1836). These efforts were patterned after the G?ttingen Magnetic Union, which also used term days and instituted a vast network of magnetic observers operating on a common plan. As with the IPYs, which cited these precedents in instituting its own term days, simultaneous observations were meant to foster both scientifi c understanding and peaceful international cooperation. FIGURE 1. Smithsonian Institution ca. 1860, home of the Meteorological Project. Source: Smithsonian Institution. 01_Fleming_pg001-012_Poles.indd 201_Fleming_pg001-012_Poles.indd 2 11/17/08 8:31:07 AM11/17/08 8:31:07 AM ADVANCING POLAR RESEARCH 3 James P. Espy (1785? 1860), the fi rst meteorologist employed by the U.S. government, captured the basic dif- ference between the lone astronomer and the needs of the gregarious meteorologist: The astronomer is, in some measure, independent of his fellow astronomer; he can wait in his observatory till the star he wishes to observe comes to his meridian; but the meteorolo- gist has his observations bounded by a very limited horizon, and can do little without the aid of numerous observers furnishing him contemporaneous observations over a wide-extended area. (Espy, 1857:40) Espy worked closely with Joseph Henry (1797? 1878), the fi rst secretary of the Smithsonian Institution, to create a meteorological network of up to 600 volunteer observ- ers, reporting monthly, that spanned the entire United States and extended internationally. Some telegraph stations also cooperated, transmitting daily weather re- ports to Washington, D.C., where the information was posted on large maps in the Smithsonian Castle (Figure 1) and at the U.S. Capitol. The Smithsonian meteorologi- cal project provided standardized instruments, uniform procedures, free publications, and a sense of scientifi c unity; it formed a ?seedbed? for the continued growth of theories rooted in data. To increase knowledge of the atmosphere, it sponsored original research on storms and climate change; to diffuse knowledge, it published and distributed free reports, instructions, and translations. It soon became the U.S. ?national center? for atmospheric research in the mid-nineteenth century, as well as a clear- inghouse for the international exchange of data (Fleming, 1990:75? 94). Nineteenth-century meteorology benefi ted from many of the leading technologies and theories available at the time, which, in turn, fueled public expectations about weather prediction. Telegraphy provided instantaneous transmission of information, at least between stations on the grid, and connected scientists and the public in a vast network of information sharing. Nineteenth-century meteorology, climatology, and other areas of geophysics were undoubtedly stimulated by telegraphic communications that enabled simultaneous observations, data sharing, and timing of phenomena such as auroras, occultations, and eclipses. The vast amounts of gathered data also encouraged scientists to experi- ment with new ways of portraying the weather and other phenomena on charts and maps (Anderson, 2006). In an effort to enhance both the understanding and prediction of weather phenomena, Yale professor Elias Loomis (1811? 1889) searched for ?the law of storms? governing storm formation and motion (Figure 2) (Fleming, 1990:77? 78, 159). He also mapped the occurrence, intensity, and fre- quency of auroras from global records, providing a pre- view of what might be accomplished by observing at high latitudes (Figure 2) (Shea and Smart, 2006). FIGURE 2. Charts by Elias Loomis, ca. 1860. (left) Trajectories of storms entering northeastern USA. Source (Fleming 1990, 159); (right) Fre- quency of aurora borealis sightings; darker band shows at least 80 auroras annually (http://www.phy6.org/Education/wloomis.html). 01_Fleming_pg001-012_Poles.indd 301_Fleming_pg001-012_Poles.indd 3 11/17/08 8:31:09 AM11/17/08 8:31:09 AM 4 SMITHSONIAN AT THE POLES / FLEMING AND SEITCHEK Public demand for weather-related information world- wide led to the establishment of many national services by the 1870s. In the United States, the Army Signal Offi ce was assigned this task and soon took the lead in international cooperation. In 1873, the U.S. proposed that all nations prepare an international series of simultaneous observa- tions to aid the study of world climatology and weather patterns. This suggestion led to the Bulletin of International Simultaneous Observations, which contained worldwide synoptic charts and summaries of observations recorded at numerous locations around the world (Figure 3) (Myer, 1874:505). Beginning in 1871, the U.S. National Weather Service issued daily forecasts, heightening public expecta- tions that weather could be known? and in some cases, prepared for? in advance. The increasing density and geo- graphic extent of information becoming available in meteo- FIGURE 3. International Synoptic Chart: Observations cover the Northern Hemisphere, except for the oceans and polar regions. Dashed lines indicate projected or interpolated data. Source: U.S. Army Signal Offi ce, Bulletin of International Meteorology, 28 January 1884. 01_Fleming_pg001-012_Poles.indd 401_Fleming_pg001-012_Poles.indd 4 11/17/08 8:31:11 AM11/17/08 8:31:11 AM ADVANCING POLAR RESEARCH 5 rology fueled hopes that the scientifi c enterprise would soon encompass the entire globe, including the polar regions. THE FIRST INTERNATIONAL POLAR YEAR The IPY-1 resulted from the ideas of the Austrian naval offi cer and polar explorer Karl Weyprecht (1838? 1881) and the organizational skills of Georg von Neumayer (1826? 1909), director of the German Hydrographical Of- fi ce, along with Heinrich Wild (1833? 1902), director of the Central Physical Observatory in St. Petersburg. Weyprecht, who co-directed the unsuccessful 1872 Austro-Hungarian North Pole Expedition, argued that decisive scientifi c re- sults could only be obtained by research stations distrib- uted over the Arctic regions and charged with the task of obtaining one year?s series of reliable meteorological and geophysical observations made with the same methods (Barr, 1983:463? 483). Weyprecht wrote in 1875: The key to many secrets of Nature . . . is certainly to be sought for near the Poles. But as long as Polar Expeditions are looked upon as merely a sort of international steeple-chase, which is primarily to confer honour upon this fl ag or the other, and their main objective is to exceed by a few miles the latitude reached by a predecessor, these mysteries will remain unsolved. (Weyprecht, 1875:33) Weyprecht formulated six principles of Arctic research: (1) Arctic exploration is of greatest importance for a knowl- edge of the laws of nature; (2) geographical discovery is of serious value only when linked to scientifi c exploration; (3) detailed Arctic topography is of secondary importance; (4) the geographic pole is of no greater importance for science than other high-latitude locations; (5) favorable locations for stations are near high-intensity phenomena; and (6) isolated series of observations are of limited value (Baker, 1982). Weyprecht?s ideas were institutionalized in 1879 with the establishment of the International Polar Commission at the German Hydrographical Offi ce, chaired by von Neumayer. The IPY-1, launched just one year after Wey- precht?s death, brought together a cast of hundreds, even- tually resulting in 11 nations placing 12 stations around the North Pole and two near the South Pole (Figure 4) (Anonymous, 1884). The scientists in IPY-1 practiced a form of coordinated Humboldtean science; that is, they emulated the exhaustive methods of the famed German scientifi c traveler Alexander von Humboldt (1769? 1859), who took precision measure- ments of natural phenomena. The expeditions set out with an ambitious agenda and tracked data for fi elds as diverse as meteorology, magnetism, glaciology, oceanography, sea ice studies, geomorphology, phytogeography, exploration, mapping, ethnography, and human geography. They made observations in conditions of hardship, hunger, extreme cold, severe gales, blinding drifting snow, and continuous darkness, with frozen instruments often coated with ice. Each station established its own identity while retain- ing its position in the greater network. The station at Point Barrow (Figure 5), established by the U.S. Army Signal Service, also served as a crucial part of the Smithsonian Institution?s natural history and ethnographic studies (Burch, this volume; Crowell, this volume; Krupnik, this volume). The Kara Sea observations were conducted on sea ice when the Norwegian steamer Varna became beset. The Varna and a Danish relief vessel Dijmphna gathered data for the entire IPY. The Varna eventually sank after being crushed by the moving ice. The signal service station at Fort Conger, Lady Franklin Bay (Figure 5), originally conceived as a base from which a U.S. expedition might reach the North Pole, met with tragedy. The expedition lost 19 of 25 men when resupply efforts failed in 1883? 1884. Yet its leader, Lt. Adolphus Greely (1844? 1935), FIGURE 4. The IPY-1 Stations (and their sponsoring countries) dur- ing 1881? 1884. Arctic (clockwise): Sagastyr (Russia, R), Kara Sea near Dicksonhafen (Netherlands, NL), Karmakuly (Russia), Sodan- kyla (Finland, FL), Bossekop (Norway, N), Cape Thordsen (Sweden, S), Jan Mayen Island (Austria, A), Godthaab (Denmark, DK), Lady Franklin Bay/Fort Conger (USA), Kingua Fjord (Germany, D), Fort Rae (Great Britain, GB), Point Barrow (USA); Southern Hemisphere: Orange Bay, Tierra del Fuego (France, F) and Royal Bay, South Geor- gia Island (Germany). Source: original graphic, after Barr, 1985. 01_Fleming_pg001-012_Poles.indd 501_Fleming_pg001-012_Poles.indd 5 11/17/08 8:31:15 AM11/17/08 8:31:15 AM 6 SMITHSONIAN AT THE POLES / FLEMING AND SEITCHEK who nearly starved to death himself, took steps to protect the instruments and data. Despite hardships and limitations, each expedition published a fi nal report accompanied by numerous scien- tifi c articles and popular accounts. The IPY-1 data were intended to enable the creation of new synoptic charts that could connect polar weather conditions to those in lower latitudes. Yet ultimately, the network of only 12 stations scattered north of 60 degrees latitude was spread too thin (cf. Wood and Overland, 2006). Alfred J. Henry (1858? 1931), chief of the meteorological records division of the U.S. Weather Bureau, observed that the ?gap between the polar stations and those of the middle latitudes [was] en- tirely too wide to span by any sort of interpolation and thus the relationship of polar weather to the weather of mid-latitudes failed of discovery.? Noted geographer Isa- iah Bowman (1878? 1950) commented in 1930, ?The fi rst polar explorers could go only so far as the state of technol- ogy and theory permitted? (Bowman, 1930: 442). Still there were modest accomplishments, for exam- ple, in expanded knowledge of the weather in the Davis Strait between Canada and Greenland and the infl uence of the Gulf Stream in northern latitudes. IPY-1 data were also used in 1924 to construct circumpolar charts for planning ?Aeroarctic,? the international airship expedition to the Russian Arctic, conducted in 1931 (Luedecke, 2004). In 2006, a systematic reanalysis and reevaluation of IPY-1 data provided insights on climate processes and points of comparison with subsequent Arctic climate patterns. While the stations showed that sea-level pressures and surface air temperatures were indeed infl uenced by large- scale hemispheric circulation patterns, in the end, the data lacked suffi cient density and the time period was too short to allow for any fundamental discoveries in meteorology or earth magnetism (Wood and Overland, 2006). TOWARD THE SECOND INTERNATIONAL POLAR YEAR As the fi ftieth anniversary of the IPY approached, the leading edge of 1930s technology, particularly avia- tion and radio, provided scientists with new capabilities for collecting data and collaborating with colleagues on projects of global scale. New theories and new organiza- tions supported the geosciences, while public expectations for weather and climate services continued to rise. The ?disciplinary? period in meteorology began in the second decade of the twentieth century, rather late compared to parallel developments in other sciences, but just in time to inform planning for IPY-2. Meteorologists in World War I were trained to analyze and issue battlefi eld weather maps; to take hourly measurements conducive to launch- ing and defending against poison gas attacks; and to col- lect data on upper-air conditions, especially winds, to help calculate the trajectories of long-range artillery shells (Bates and Fuller, 1986:15? 19; Fuller, 1990:9? 15). By us- ing pilot balloons with theodolite trackers and electrical timers, observers could track the winds and atmospheric conditions aloft. FIGURE 5. U.S. IPY-1 stations at (left) Point Barrow, Alaska, and (right) Fort Conger, Lady Franklin Bay, Canada. Source: Wood and Overland 2007. 01_Fleming_pg001-012_Poles.indd 601_Fleming_pg001-012_Poles.indd 6 11/17/08 8:31:17 AM11/17/08 8:31:17 AM ADVANCING POLAR RESEARCH 7 Meteorologists also provided critical support for avia- tion and benefi ted, especially after the war, by data col- lection from instrumented aircraft, using wing-mounted aerometeorographs that continuously recorded atmo- spheric data. By 1920, the Bergen school of meteorology in Norway had fi rmly established the principles of air- mass analysis. Of greatest relevance to polar meteorology are the massive domes of clear cold air called continental arctic air masses that sweep across Canada and Siberia, dramatically infl uencing the weather in lower latitudes. The so-called polar air masses? both continental and maritime? are also signifi cant weather-makers, although they originate below the Arctic Circle. Also, using newly available information on the vertical structure of the at- mosphere, Bergen meteorologists identifi ed inclined sur- faces of discontinuity separating two distinct air masses, most notably the polar front that spawns many severe winter storms (Figure 6). These conceptual models, com- bined with objective techniques of weather map analysis and the hope of someday solving the complex equations of atmospheric motion governing storm dynamics, breathed new theoretical life into what had been a largely empirical and applied science (Friedman, 1982). Radio technology also provided new scientifi c capa- bilities. Since magnetic disturbances and auroral displays interfered with radio transmission and telephone wires, radio equipment could be used to detect these phenomena and measure their strength. Radio also provided precise time signals to coordinate simultaneous measurements and communication links that allowed the polar stations to stay in touch with each other and with supporters in lower latitudes (Figure 7, left). Balloon-borne radiosondes used miniature transmitters to send pressure, temperature, and humidity to earth from altitudes as high as 10 kilome- ters. (Figure 7, right). Special sondes were outfi tted to take measurements of cosmic rays, ultraviolet light, ozone, and other data previously gathered with self-registering bal- loonsondes; this provided a signifi cant advantage since it was almost impossible to recover meteorographs launched in remote polar areas (DuBois et al., 2002). Public expectations about climate and weather broad- ened as radio broadcasts of weather conditions became commonplace. As weather broadcasts increased, so did the number of people employed in weather reporting. Ra- dio in the mid-1920s created the ?weather personality,? which became an established role at many stations. One of the fi rst weather personalities was E. B. Rideout at sta- tion WEEI that started broadcasting from Boston in 1924 (Leep, 1996). Early commercial airlines also benefi ted from the im- proved weather data. An airways weather service provided valuable information to pilots and dispatchers in support of commercial aviation, which navigated by landmarks and instrument readings. In 1927, based on signifi cant technological advances, new theories of dynamic meteorology, and rising public expectations, the German meteorologist Johannes Georgi (1888? 1972) raised the issue of a possible second Interna- tional Polar Year. Two years later, the International Con- ference of Directors of Meteorological Services at Copen- hagen approved the following resolution: Magnetic, auroral and meteorological observations at a network of stations in the Arctic and Antarctic would materi- ally advance present knowledge and understanding [of these phenomena] not only within polar regions but in general ... this increased knowledge will be of practical application to problems connected with terrestrial magnetism, marine and aerial naviga- tion, wireless telegraphy and weather forecasting. (C. Luedecke, 2006, cited with author?s permission) IPY-2 was held in 1932? 1933, the fi ftieth anniversary of IPY-1. Although a worldwide economic depression lim- ited participation, some 40 nations sent scientifi c teams to reoccupy the original stations and open new ones. Re- search programs were conducted in meteorology, terres- trial magnetism, atmospheric electricity, auroral physics, and aerology using the newest technologies of radio com- munication. As in previous fi eld programs, certain peri- ods, now called ?international days,? were designated for intensive, around-the-clock observations. Even in that era, scientists detected signs of Arctic warming. FIGURE 6. Norwegian model of the polar front through a series of cyclones (Bjerknes and Solberg, 1922). 01_Fleming_pg001-012_Poles.indd 701_Fleming_pg001-012_Poles.indd 7 11/17/08 8:31:19 AM11/17/08 8:31:19 AM 8 SMITHSONIAN AT THE POLES / FLEMING AND SEITCHEK Also, the Second Byrd Antarctic Expedition of 1933? 1935? that coincided with, and expanded beyond, IPY- 2? brought new focus to Antarctica. It established a year-round meteorological station on the Ross Ice Shelf and captured public attention through live weekly radio broadcasts. Several additional IPY-2 stations in low lati- tudes added to the worldwide nature of the effort. The IPY-2 benefi ted from a sense of interconnected- ness stimulated by the International Union of Geodesy and Geophysics (IUGG), a nongovernmental, scientifi c organi- zation founded in 1919 to promote both disciplinary ad- vances and the ultimate unity of the planetary sciences. The polar front theory also provided focus. As Isaiah Bowman remarked, IPY-2 meteorologists, especially those trained in Norwegian methods, were ?inspired by a profound curiosity as to the suspected infl uence of weather condi- tions in high latitudes upon (or interaction with) those of the temperate regions as well as the tropics? (Bowman, 1930:442). The IPY-2 accomplishments included simulta- neous measurements at multiple stations; higher temporal and spatial resolution, including the vertical; and new in- strumentation such as radiosondes, ionosondes, rapid-run magnetometers; and accurate timing of global current pat- terns for magnetic storms. Reporting included the launch of the Polar Record in 1931, an international journal on polar research published in Cambridge, UK, numerous sci- entifi c papers, articles, personal accounts, data archives, and a comprehensive IPY-2 bibliography published in 1951, just in time for planning the IGY (Laursen, 1951). However, an expected summary publication for IPY-2 was not produced until 1959 (Laursen, 1959), and a part of the IPY-2 instrumental data was lost at its major inter- national depository in Copenhagen, presumably during World War II. THE DAWN OF THE INTERNATIONAL GEOPHYSICAL YEAR World War II swelled the ranks of practicing meteo- rologists and introduced new technologies originally de- veloped for the military. Rockets provided a new means for accessing upper levels of the atmosphere, broadening the scope of data collection. Not only could instruments be sent high into and even beyond the atmosphere to take measurements, but also cameras could travel to new heights to send back images of the earth. Electronic com- puters, designed to crack codes, calculate shell trajecto- ries, and estimate bomb yields, were applied to problems of geophysical modeling, while radar enabled scientists to FIGURE 7. (left) W. C. Brown operating radio equipment at Simavik, Norway, during IPY-2 (http://www.wdc.rl.ac.uk/ionosondes/history/IPY. html); (right) John Rea, left, and Stuart McVeigh launching a radiosonde at Chesterfi eld Inlet, Hudson Bay, Canada, in winter. Source: Univer- sity of Saskatchewan Archives. 01_Fleming_pg001-012_Poles.indd 801_Fleming_pg001-012_Poles.indd 8 11/17/08 8:31:20 AM11/17/08 8:31:20 AM ADVANCING POLAR RESEARCH 9 visualize weather patterns remotely. An important aspect of this new technological age was atmospheric nuclear testing, which injected radionuclide ?tracers? into the environment. Meteorology, climatology, and aeronomy? ?the atmospheric sciences?? benefi ted intellectually from an infl ux of new talent from fi elds such as mathematics, physics, chemistry, and engineering. The same technology that provided remote-imaging capabilities for scientists was also used in broadcasting to reach the general public. While long-range weather fore- casts and even weather control were distinct possibilities, the public was growing apprehensive about the meteoro- logical effects of atmospheric nuclear testing and increas- ingly visible levels of smoke and smog. Scientists were in- creasingly interested in the interconnected workings of the global environment, while military planners sought new geophysical capabilities (Fleming, 2000). American physicist Lloyd Berkner (1905? 1967) sug- gested that IPY-3 take place 25 years after IPY-2. His col- league, Sidney Chapman, who suggested that the event be called the International Geophysical Year, served as president of Comit? Sp?cial de l?Ann?e G?ophysique In- ternationale (CSAGI), which coordinated the effort inter- nationally. In his presidential remarks, Chapman (1959b) emphasized the earth?s fl uid envelope and the continuing need for widespread simultaneous observations: The main aim [of the IGY] is to learn more about the fl uid envelope of our planet? the atmosphere and oceans? over all the earth and at all heights and depths. The atmosphere, especially at its upper levels, is much affected by disturbances on the sun; hence this also will be observed more closely and continuously than hitherto. Weather, the ionosphere, the earth?s magnetism, the polar lights, cosmic rays, glaciers all over the world, the size and form of the earth, natural and man-made radioactivity in the air and the seas, [and] earthquake waves in remote places will be among the subjects studied. These researches demand wide- spread simultaneous observation. The IGY logo emphasized the infl uence of the sun on the earth, scientifi c focus on Antarctica, and the hope that geophysical satellites would soon be placed in orbit (Fig- ure 8). The breadth of the program was certainly made possible by new technological developments in transpor- tation, communication, and remote sensing. Teams of observers equipped with the latest scientifi c instruments were deployed around the globe? some to the ends of the earth in polar regions, on high mountaintops, and at sea? to study earth processes. The effort in Antarctica alone involved hundreds of people in logistically complex and expensive expeditions. While Earth-orbiting satellites were in their infancy, Explorer 1 and Explorer 3 brought immediate geophysical results that fundamentally altered our understanding of the planet? the discovery of the Van Allen radiation belts (National Research Council, 2007). The IGY?s 18 months of comprehensive global research resulted in other accomplishments as well, including the charting of ocean depths and currents, an in-depth study of Antarctic ice sheets, and, notably, the beginnings of global CO 2 monitoring efforts. The IGY captured scien- tifi c center stage at the time and generated many technical and popular publications. Its organizers, recognizing that the international interchange of geophysical data was ?the immediate and specifi c end of its vast scientifi c program,? also made careful provisions for its preservation in the World Data Centers (Odishaw, 1962). The IGY was actually the twenty-fi fth anniversary of IPY-2. Had there been a full fi ftieth anniversary, it would have occurred in 1982? 1983, well into the era of Earth satellite observations that by then were providing com- plete coverage of many global atmospheric processes. In the late 1970s, the Global Atmospheric Research Program (GARP) was gearing up for the Global Weather Experiment (GWE)? at the time the largest fully international scientifi c experiment ever undertaken? linking in situ and satellite data to computer modeling in an attempt to improve op- erational forecasting, determine the ultimate range of nu- merical weather prediction, and develop a scientifi c basis for climate modeling and prediction. In this experiment, FIGURE 8. The IGY logo adopted in 1955 and used on IGY instruments and publications. Source: U.S. National Academy of Sciences. 01_Fleming_pg001-012_Poles.indd 901_Fleming_pg001-012_Poles.indd 9 11/17/08 8:31:22 AM11/17/08 8:31:22 AM 10 SMITHSONIAN AT THE POLES / FLEMING AND SEITCHEK worldwide surface and upper-air observations from satel- lites, ships, land stations, aircraft, and balloons were com- bined with global coverage provided by fi ve geostationary satellites operated by the United States, Russia, Japan, and the European Space Agency (ESA) (Figure 9). Polar orbit- ing satellites covered the rest of the globe. This experiment was followed by the World Climate Research Programme (WCRP), which began in 1980. Today, it can be said that the global observing system provides the equivalent of a GWE of data every day (National Research Council, 2007). The challenge today lies in analyzing, assimilating, and ar- chiving the massive fl ows of data. CONCLUSIONS? TOWARD IPY 2007? 2008 As we enter the International Polar Year of 2007? 2008, today?s scientists are heirs to a grand research tradi- tion. With more than 60 nations participating, the current IPY has a broad interdisciplinary focus on environmental change. Six scientifi c themes provide a framework for IPY 2007? 2008 (Allison et al., 2007:13): 1. Status: to determine the present environmental status of the polar regions; 2. Change: to quantify and understand past and present natural environmental and social change in the polar regions and to improve projections of future change; 3. Global linkages: to advance understanding on all scales of the links and interactions between polar regions and the rest of the globe, and of the processes controlling these; 4. New frontiers: to investigate the frontiers of science in the polar regions; 5. Vantage point: to use the unique vantage point of the polar regions to develop and enhance observatories from the interior of the earth to the sun and the cosmos beyond; and 6. The human dimension: to investigate the cultural, his- torical, and social processes that shape the sustainabil- ity of circumpolar human societies and to identify their unique contributions to global cultural diversity and citizenship. The current IPY is supported by the latest technolo- gies including computer models of ice sheet dynamics, ice cores reaching all the way to bedrock, and advanced sur- face, airborne, and satellite sensors that measure ice thick- ness, surface elevation, mass balance, and subsurface con- ditions. Recently, scientists have discovered rapid changes FIGURE 9. During the Global Weather Experiment of 1979, fi ve international geostationary satellites supported global mid-latitude observa- tions of cloud-tracked winds. (GMS H11005 Geostationary Meteorological Satellite; GOES H11005 Goestationary Operational Environmental Satellites; METEOSAT, ESA H11005 a meteorological satellite (European Space Agency); GOMS H11005 Geostationary Operational Meteorological Satellite) 01_Fleming_pg001-012_Poles.indd 1001_Fleming_pg001-012_Poles.indd 10 11/17/08 8:31:23 AM11/17/08 8:31:23 AM ADVANCING POLAR RESEARCH 11 in ice sheets. The latest collapse of the Larsen B ice shelf in Antarctica in 2002, captured only because of frequent coverage by satellite imagery, illustrated this dynamic on astonishingly short time scales. These new ?bits? of knowledge carry weighty implications: The rapid transfer of ice from the continental ice sheets to the sea could re- sult in a signifi cant rise of sea level. Because of the global implications of the changes at the poles for ecosystems and human communities everywhere, IPY 2007? 2008 science aims to reach a wide audience, train a new generation of polar researchers, and galvanize public opinion through the associated education, outreach, and communication efforts. The 125-year history of polar scientifi c quests, like the much longer history of cooperative observations, involves scientifi c research questions, technological (including logis- tic) capabilities, and public perceptions. Since 1882? 1883, all IPY ventures have been about science done in extreme conditions, fruitful international cooperation, arctic air masses and polar fronts, melting ice caps, and rising sea levels. That is, the IPY studies are essentially ?about us.? Each successive IPY has been based upon the technologi- cal innovations of its era and the leading scientifi c theories and hypotheses developed by its time. In 1881 at the start of IPY-1, the Russian meteorolo- gist Heinrich Wild observed that ?the good and favor- able idea of Weyprecht . . . has survived the calamities of war, the discords of nations, the obstacles of jealous people, and the death of the author? (Baker, 1982:284). Today, in spite of intervening world wars and numerous discords and obstacles, this statement still rings true. The International Polar Year 2007? 2008 is solidly grounded in scientifi c cooperative efforts for the increase and diffu- sion of knowledge. Its goal, like those of its predecessors, is to advance research and communicate its wonders. It also aims to preserve the habitability of the planet. Joseph Henry would be pleased. LITERATURE CITED Allison, I., M. B?land, K. Alverson, R. Bell, D. Carlson, K. Dannel, C. Ellis-Evans, E. Fahrbach, E. Fanta, Y. Fujii, G. Glasser, L. Gold- farb, G. Hovelsrud, J. Huber, V. Kotlyakov, I. Krupnik, J. Lopez- Martinez, T. Mohr, D. Qin, V. Rachold, C. Rapley, O. Rogne, E. Sarukhanian, C. Summerhayes, and C. Xiao. 2007. The Scope of Science for the International Polar Year 2007? 2008. World Meteo- rological Organization, Technical Document 1364. Geneva: World Meteorological Organization. Anderson, K. 2006. ?Mapping Meteorology.? In Intimate Universality: Local and Global Themes in the History of Weather and Climate, ed. J. R. Fleming, V. Jankovic, and D. R. Coen, pp. 69? 92. Saga- more Beach, Mass.: Science History Publications. Anonymous. 1884. The International Polar Stations. Science, 4:370? 372. Baker, F. W. G. 1982. The First International Polar Year, 1882? 1882. Polar Record, 21:275? 285. Barr, W. 1983. Geographical Aspects of the First International Polar Year, 1882? 1883. Annals of the Association of American Geogra- phers, 73: 463? 483. ??? . 1985. The Expeditions of the First International Polar year, 1882? 1883. The Arctic Institute of North America, Technical Pa- per 29. Calgary, Alberta, Canada: University of Calgary. Bates, C. C., and J. F. Fuller. 1996. America?s Weather Warriors, 1814? 1985. College Station: Texas A&M University Press. Bjerknes, J., and H. Solberg, 1922. Life Cycle of Cyclones and the Po- lar Front Theory of Atmospheric Circulation, Geofysiske Publika- tioner 3(1): 3? 18. Bowman, I. 1930. Polar Exploration. Science, n.s., 72: 439? 449. Burch, Ernest S., Jr. 2009. ?Smithsonian Contributions to Alaskan Eth- nography: The IPY Expedition to Barrow, 1881? 1883.? In Smith- sonian at the Poles: Contributions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 89? 98. Washington, D.C.: Smithsonian Institution Scholarly Press. Cassidy, D. C. 1985. Meteorology in Mannheim: The Palatine Meteoro- logical Society, 1780? 1795. Sudhoffs Archiv: Zeitschrift f?r Wis- senschaftsgeschichte, 69: 8? 25. Chapman, S. 1959a. IGY: Year of Discovery. Ann Arbor: University of Michigan Press. ??? . 1959b. Presidential Address, 28 January 1957. Annals of the International Geophysical Year, 1: 3? 5. Crowell, Aron L. 2009. ?The Art of I?upiaq Whaling: Elders? Interpre- tations of International Polar Year Ethnological Collections.? In Smithsonian at the Poles: Contributions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 99? 114. Washington, D.C.: Smithsonian Institution Scholarly Press. DuBois, J. L., R. P. Multhauf, and C. A. Ziegler. 2002. The Invention and Development of the Radiosonde with a Catalog of Upper- Atmospheric Telemetering Probes in the National Museum of American History. Smithsonian Studies in History and Technology, No. 53. Washington, D.C.: Smithsonian Institution Press. Espy, J. P. 1857. Fourth Meteorological Report. U.S. Senate, Ex. Doc. 65, 34th Cong., 3rd sess. Washington, D.C. Fleming, J. R. 1990. Meteorology in America, 1800? 1870. Baltimore: Johns Hopkins University Press. ??? , ed. 2000. Geophysics and the Military. Historical Studies in the Physical and Biological Sciences 30(2). [Special issue] Friedman, R. M. 1982. Constituting the Polar Front, 1919? 1920. Isis, 74: 343? 362. Fuller, J. F. 1990. Thor?s Legions. Boston: American Meteorological Society. Heathcote, N., and A. Armitage. 1959. The First International Polar Year (1882? 1883). Annals of the International Geophysical Year, 1: 6? 100. Herschel, J. 1836. Report of the Meteorological Committee of the South African Literary and Scientifi c Institution. Edinburgh New Philo- sophical Journal, 21: 239? 246. Krupnik, Igor. 2009. ??The Way We See It Coming?: Building the Legacy of Indigenous Observations in IPY 2007? 2008.? In Smithsonian at the Poles: Contributions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 129? 142. Washington, D.C.: Smithsonian Institution Scholarly Press. Laursen, V. 1951. Bibliography for the Second International Polar Year 1932? 3. Copenhagen, Denmark: Horsholm Bogtrykkeri. ??? . 1959. The Second International Polar Year. Annals of the Inter- national Geophysical Year, 1: 211? 234. Leep, R. 1996. ?The American Meteorological Society and the Develop- ment of Broadcast Meteorology.? In Historical Essays on Meteorol- ogy, 1919? 1995, ed. J. R. Fleming, pp. 481? 507. Boston: American Meteorological Society. 01_Fleming_pg001-012_Poles.indd 1101_Fleming_pg001-012_Poles.indd 11 11/17/08 8:31:41 AM11/17/08 8:31:41 AM 12 SMITHSONIAN AT THE POLES / FLEMING AND SEITCHEK Luedecke, C. 2004. The First International Polar Year (1882? 83): A Big Science Experiment with Small Science Equipment. History of Meteorology, 1: 55? 64. ???. 2006. Changing Trends in Polar Research as Refl ected in the History of the International Polar Years. Unpublished manuscript. Institute for History of Science and Technology, University of Hamburg, Germany. Myer, A. J. 1874. Annual Report of the U.S. Army Signal Offi ce, 1874. Washington, D.C. National Research Council. 2007. Earth Observations from Space: The First 50 Years of Scientifi c Achievements. Washington, D.C.: Na- tional Academies Press. Odishaw, H. 1962. What Shall We Save in the Geophysical Sciences? Isis, 53: 80? 86. Shea, M. A., and D. F. Smart. 2006. Compendium of the eight articles on the ?Carrington Event? attributed to or written by Elias Loomis in the American Journal of Science, 1859? 1861. Advances in Space Research, 38(2): 313? 385. Societas Meteorologica Palatina. 1783? 1795. Ephemerides, 12 vols. Weyprecht, K. 1875. Scientifi c Work on the Second Austro-Hungarian Polar Expedition, 1872? 4. Royal Geographical Society Journal, 45:19? 33. Wood, K. R., and J. E. Overland. 2006. Climate Lessons from the First International Polar Year. Bulletin of the American Meteorological Society, 87: 1685? 1697. ???. 2007. Documentary Image Collection from the First Inter- national Polar Year, 1881? 1884. http://www .arctic .noaa .gov/ aro/ ipy-1/ Frontpage.htm (accessed 2 November 2007). 01_Fleming_pg001-012_Poles.indd 1201_Fleming_pg001-012_Poles.indd 12 11/17/08 8:31:42 AM11/17/08 8:31:42 AM ABSTRACT. The fi rst International Polar Year (IPY) of 1882? 1883 came at the end of a half-century of efforts at collaborative and/or cooperative research among the scientifi c communities of Europe and the United States. These efforts included the Magnetic Cru- sade, a cooperative endeavor to solve fundamental questions in terrestrial magnetism; a variety of plans for international cooperation in the gathering of meteorological data; the observations of the transits of Venus; and the establishment of the Smithsonian?s interna- tional network to alert astronomers of new phenomena. It was also a half century when scientifi c exploration of the polar regions was still problematic in terms of the safety and survival of the investigator. This paper will look at scientifi c cooperation and earlier Polar research as the background for the fi rst IPY, with special emphasis on the leadership role taken by the Smithsonian Institution. INTRODUCTION The fi rst International Polar Year (IPY), which included 14 expeditions spon- sored by 11 countries (12 expeditions to the North Polar Region, 2 to the South Polar Region), was a landmark event in the history of polar science. During the half-century leading up to the coordinated research efforts of 1882? 1883, scien- tifi c research in the Polar Regions had been very problematic. Survival, let alone the successful completion of observations, was uncertain. The use of trained specialists was a rarity. Instead, research was usually conducted as a sideline to the primary objectives or mission of the expedition, which were geographi- cal discovery, by a scientifi cally inclined explorer, military offi cer, or physician who made observations or collected specimens on a limited basis. Attempting to reach higher latitudes was an end in itself, a form of international competition, independent of any scientifi c return (Barr, 1983: 464). The catalyst for the transformation from competition and exploration to cooperation and scientifi c research was Karl Weyprecht, the Austrian explorer who fi rst suggested the IPY. It was Weyprecht?s ?drive, ambition, and connec- tions? which were essential in bring the idea of an international, cooperative attack on the problems of polar science to fruition, although he died in 1881, before the IPY was offi cially launched (Barr, 1983: 464). Marc Rothenberg, Historian, National Science Foundation, 4201 Wilson Boulevard, Arlington, VA 22230, USA (mrothenb@nsf.gov). Accepted 29 May 2008. Cooperation at the Poles? Placing the First International Polar Year in the Context of Nineteenth-Century Scientifi c Exploration and Collaboration Marc Rothenberg 02_Rothenberg_pg013-000_Poles.in13 1302_Rothenberg_pg013-000_Poles.in13 13 11/17/08 8:36:48 AM11/17/08 8:36:48 AM 14 SMITHSONIAN AT THE POLES / ROTHENBERG It is important, however, not to claim too much for the fi rst IPY. It did not launch science on an entirely new path of international cooperation. That was already a well-trod path by the last quarter of the nineteenth-century, and many of the programs of the Smithsonian Institution, for example, incorporated some aspect of international coop- eration. If it proved ?that international scientifi c ventures were possible on a large scale,? as C. J. Taylor (1981: 376) contended, it was just one of many proofs. If it demon- strated that scientists could cooperate in spite of national differences at a time that when international relations were fraught with danger (Budd, 2001: 50? 51), so too did scientists in a variety of other disciplines cooperate during this era in order to further research on a number of differ- ent scientifi c questions. The IPY was organized at the end of a half century marked by efforts at collaborative research or other forms of cooperation in the physical sciences among and be- tween the scientifi c communities of Europe and the United States. These efforts included the Magnetic Crusade, a collaborative endeavor to solve fundamental questions in terrestrial magnetism; a variety of plans for international cooperation in the gathering of meteorological data; the establishment of the Smithsonian?s international network to alert astronomers of new phenomena; and the many expeditions sent out throughout the world to observe the transits of Venus. The level of cooperation ranged from simply improving communication among scientists to es- tablishing common standards for recording observations. This urge to cooperate across national boundaries in the nineteenth century was not limited to the world of sci- ence. It was an integral part of the Victorian-era Euro- American society. Perhaps this urge was most clearly expressed through the organization of international con- gresses. For example, no less than 32 international con- gresses met in conjunction with the 1878 Exposition at Paris. The various agendas included cooperation, coor- dination, standardization, exchange of information, best methods for the gathering of statistics, and efforts at com- mon solutions for common problems. The congresses ranged in subject area from legal issues, such as interna- tional copyright, patent rights, and legal medicine to social issues, such as prevention of cruelty to animals, the treat- ment of alcoholism, guidelines for military ambulance ser- vice, and aid to the blind and deaf. Science was not left out. Among the scientifi c fi elds to hold congresses in Paris that year were geometry, anthropology, ethnography, bot- any, geology, and meteorology. Included on the agendas of the scientifi c congresses were such issues as simultaneity of observations and uniform nomenclatures. There was also a congress to discuss the possibility of the adoption of a uniform system of weight, measures, and coinage (United States, 1880: 1: 455? 464). In this paper, I will briefl y summarize the various nineteenth-century efforts at international collaboration and cooperation in science, with particular attention to the role played by the Smithsonian Institution and its leader, Joseph Henry. From this discussion, it should be evident that a proposal for international cooperation to solve a scientifi c question, such as the proposal for the fi rst IPY, would not appear to be a startling new idea to European or American physical and earth scientists in the 1870s or 1880s. In fact, just the opposite was true; by the last quar- ter of the nineteenth century, efforts at cooperation were the norm, not the exception. The story is not one of inevi- table success. Sometimes the efforts at cooperation failed. The general movement of the international physical science community, however, was toward better communication and coordination. Rather than look at the IPY as a new beginning, it is more accurate, I believe, to look at it as a culmination. What occurred with the fi rst IPY was not a revolution in international science, but the transformation of polar science; it began to more closely resemble the norm in international science. MAGNETIC CRUSADE The fi rst great international effort at coordinating physical science research in the nineteenth century was the Magnetic Crusade, which focused on the international gathering of terrestrial magnetic observations (Cawood, 1977; 1979). The roots of the Magnetic Crusade lay in the appreciation by early nineteenth-century scientists that the variations of the earth?s magnetic fi eld were ex- tremely complex. Driven by both the desire to understand geomagnetic activity and the hope of creating a practical system of navigation through geomagnetic observations, observers created an informal system of contacts ?to pro- vide a degree of order in the sometimes spasmodic and rather uncoordinated work? and of course, to exchange information? (Cawood, 1979: 496). Although Alexander von Humboldt put together a loose association of magnetic observatories linked through Paris, which had been the center of terrestrial magnetic ob- servations early in the century, a more important, and more formal, system was organized in the German-speaking world. Carl Friedrich Gauss and Wilhelm Weber founded a system in 1834 under the name of the G?ttingen Magne- tische Verein. Inspired by the work on the Continent, the 02_Rothenberg_pg013-000_Poles.in14 1402_Rothenberg_pg013-000_Poles.in14 14 11/17/08 8:36:49 AM11/17/08 8:36:49 AM COOPERATION AT THE POLES? 15 British Association for the Advancement of Science agreed, in 1838, to establish its own system. Led by Samuel Hunt Christie, John Herschel, Humphrey Lloyd, and Edward Sabine, the British Association system consisted of 10 ob- servatories, with coverage expanded to include the Brit- ish colonies and India (with the cooperation of the East India Company). The British system coordinated with 23 other observatories scattered in the Russian Empire, Asia, North America, North Africa, and Europe, all of whom were funded by their respective governments, except for those in the United States funded by academic institutions (Girard College and Harvard University). Also part of this effort was a British naval expedition to make observations in Antarctica, led by James Clark Ross (1839? 1843). There were some limitations to the international cooperation. Although the British system synchronized ob- servations using G?ttingen Mean Time, as suggested by the G?ttingen Magnetische Verein, so that data could be com- pared, there was no formal collaboration. The Paris Obser- vatory acted independently of its other European counter- parts. Nonetheless, by the time the Crusade formally ended in 1848, there was a fi rmly established network of magnetic observatories in Europe, throughout the British Empire, and in the United States that continued to make observa- tions and exchange data. Other observatories later joined in the cooperative venture, including that of the Smithso- nian Institution (Rhees, 1859: 27? 29). Most importantly, as Cawood argued, the Magnetic Crusade demonstrated ?that large-scale operations could be organized and carried through? (1979: 516). Even Taylor, who argued for the sig- nifi cance of the IPY in the demonstration of the possibilities of large-scale international cooperation, admitted that the Magnetic Crusade ?provided many precedents for subse- quent global scientifi c endeavours? (1981: 370). METEOROLOGICAL COOPERATION Weather does not respect political boundaries, and many meteorologists realized the need for cooperation. German meteorologists took the lead, with such orga- nizations as the S?ddeutsche Meteorologische Verein (1841), the K?nglich Preussische Meteorologische In- stitut (1847), and the Norddeutsche Seewarte (1872). These organizations had relatively limited geographical coverage, however, and were international only because of the political fragmentation of the German scientifi c community (Fleming, 1990: 165? 166). A more signifi cant international approach to meteoro- logical observations took place in the United States. Per- haps not coincidently, it was fi rst directed by a physicist who was an active geomagnetic observer, who had coop- erated with the Magnetic Crusade, and was aware of the rewards and challenges of international cooperation. Not only had Joseph Henry received practical advice on observ- ing from Edward Sabine while in England in 1837 (Rein- gold et al., 1979: 312? 313), but he also ?had conversation with Mr[.] Christie on the subject of establishing mag- netic observator[i]es to cooperate with those established by Humboldt? (Reingold et al., 1979: 303). Joseph Henry became the fi rst secretary of the Smithsonian Institution in 1846 and established a program that placed an emphasis on the coordination of large-scale research projects, argu- ing that there were no other institutions in the United States equipped to do so. The fi rst such project Henry embraced was the development what Elias Loomis, one of Henry?s consultants in meteorology, characterized as ?a grand me- teorological crusade? for collecting meteorological obser- vations (Smithsonian Institution, 1848: 207). The system devised by Henry had two distinct but interrelated components, both requiring cooperation. The fi rst was a system of observers who? using standard apparatus, techniques, and forms to the greatest extent possible? maintained monthly logs of weather condi- tions that were sent to the Smithsonian for reduction. These logs were used to understand climate and weather tendencies over the long term. From the onset, it was recognized that ?to give this system its greatest effi ciency, the co- operation of the British government and of the Hudson?s Bay Company [in Canada] is absolutely indis- pensable? (Smithsonian, 1848: 207). Both the British gov- ernment and the private Hudson?s Bay Company quickly agreed to cooperate (Fleming, 1990: 123). The program soon expanded throughout North and Central America. Observers were recruited in Bermuda, Mexico, all the Central American countries, and throughout the West In- dies, frequently drawing, in the latter two regions, upon Americans residing overseas (Smithsonian Institution, 1872: 68? 69). The second component was the use of the telegraph to forward data on weather in real time to the Smithsonian, allowing, in the late 1850s, for the publica- tion of the fi rst scientifi cally based weather forecasts in newspapers and the fi rst publicly posted weather maps. These forecasts were based on the conclusions drawn from the monthly data logs. Unlike the data gathering, the forecasting only lasted a few years and ended before the dream of making it international was accomplished. Among the obstacles it ran into was the realization by the commercial telegraph companies that weather data was a valuable commercial commodity; the companies 02_Rothenberg_pg013-000_Poles.in15 1502_Rothenberg_pg013-000_Poles.in15 15 11/17/08 8:36:49 AM11/17/08 8:36:49 AM 16 SMITHSONIAN AT THE POLES / ROTHENBERG wanted to charge for the use of the lines (Fleming, 1990: 145; Rothenberg et al., 2007: 102). Henry?s international system worked in part because there were no government meteorologists involved who felt the need to protect their own national systems. In- stead, Henry was relying on an international network of independent observers. Two efforts bracketing Henry?s establishment of the Smithsonian network demonstrated that meteorology was not yet ready for extended interna- tional cooperation. In 1845, an international meeting of scientists inter- ested in terrestrial magnetism and meteorology was held in conjunction with the meeting of the British Association for the Advancement of Science. Efforts to establish some sort of coordination of meteorological observations, akin to the Magnetic Crusade, ran into a very serious obstacle. The government meteorologists of the various European nations had too much invested in their own systems to lay them aside for some common system. As Edward Sabine remembered two decades later (1866: 30), the government meteorologists ?manifested so marked a disposition . . . to adhere to their respective arrangements in regard to instruments, times of observation, and modes of publica- tion,? as to make it clear the time for a uniform system ?had not then arrived.? Another effort came a few years later. Matthew Fon- taine Maury, a naval offi cer, oceanographer, and director of the Naval Observatory (Williams, 1963) was a keen student of meteorology. For example, he had independently recog- nized the possibilities presented for weather forecasting by the telegraph almost as early as Henry had (Fleming, 1990: 109). In 1851, a request from the British government to the United States government on behalf of the Royal Engineers, who were conducting meteorological observations through- out the empire, ended up being forwarded to Maury for a response. The Royal Engineers had suggested the need to establish a uniform system of recording meteorological data. Maury attempted to expand this request into a broad international cooperative venture covering both nautical and terrestrial meteorology. What this venture demon- strated was that the European meteorological community was still not yet ready for such a bold stroke. Although Maury did manage to organize an 1853 meeting in Brussels to which 10 nations sent representatives, the roadblocks to international exchange of information, let alone real coop- eration, were still huge. The sole major accomplishment of the meeting was the agreement that nations that did not use centigrade as the standard scale for temperature would add that scale to the standard thermometer (Fleming, 1990: 107? 109; Anderson, 2005: 245). After 1870, a new player in American and interna- tional meteorology appeared. Albert J. Myer, the com- mander of the United States Army Signal Corps, seized on the transmission of storm information as a worthy re- sponsibility for a military organization facing budget cuts. Eventually the Smithsonian transferred its system to the Signal Corps (Hawes, 1966). Myer?s organization came to the forefront of American meteorology just when the international community was be- coming more open to the possibilities of broad cooperation. At the 1872 meeting of the Gesellschaft deutscher Natur- forscher und ?rzte in Leipzig, meteorologists called for an international gathering to further standardization and cooperation for terrestrial observations. The result was the 1873 congress in Vienna, which ultimately attracted repre- sentatives from 20 nations. At the Vienna Congress, Myer?s proposal for international simultaneous observations was agreed to, leading to the Bulletin of International Simulta- neous Observations, fi rst published by the Signal Offi ce in 1875 (Hawes, 1966). There were, however, still obstacles to be overcome, such as the continuing confl ict between the metric and English systems of measurement (Anderson, 2005: 246). Even so, the discussions had begun (Luedecke, 2004) and, with the second international meteorological congress of 1879, held in Rome, ?a pattern of voluntary cooperation between meteorologists on inter national prob- lems? had been established which bypassed the national meteorological organizations (Weiss, 1975: 809). COOPERATION IN ASTRONOMY Henry and the Smithsonian were involved in other in- ternational cooperative efforts, for example in astronomical communication. As the quantity and quality of telescopes increased in the nineteenth century, so did the number of comets and asteroids discovered. C. H. F. Peters, professor of astronomy at Hamilton College in upstate New York, a prolifi c discoverer of asteroids, was aware of the impor- tance of the dissemination of observations to other astron- omers to aid in the calculation of orbits (or even the relo- cation of the object). Because he was also German-born and educated, he was in closer touch with his colleagues on the European continent than most of his colleagues in American observatories (Rothenberg, 1999) He wrote to Henry in January 1872, suggesting a system of communi- cating discoveries among the world?s astronomers using the Atlantic cable and the land telegraph systems of the U.S. and Europe. Peters?s system would be modeled after the Smithsonian international exchange system for publi- 02_Rothenberg_pg013-000_Poles.in16 1602_Rothenberg_pg013-000_Poles.in16 16 11/17/08 8:36:50 AM11/17/08 8:36:50 AM COOPERATION AT THE POLES? 17 cations, in which the Smithsonian served as the intermedi- ary between American scientists and scientifi c institutions seeking to distribute their publications throughout the world, and their foreign counterparts seeking to distribute publications in the United States. In the case of astronomy, the Smithsonian would serve as the American node, receiv- ing announcements of discoveries and distributing them to two proposed European nodes? the observatories at Leipzig and Vienna? and vice versa. Given Henry?s well- known inclination to support international cooperation, Peters expressed his optimism that the Smithsonian would be willing to pick up the cost of trans-Atlantic telegraph transmission (Rothenberg et al., 2007: 447) Henry, responding as Peters had anticipated, immedi- ately began seeking support for Peters?s plan. It took eigh- teen months for Peters?s proposal to be fully implemented, in part because Henry wanted to avoid having science pay for the use of the telegraph. Within a year, Henry had se- cured the support of Cyrus Field, the father of the Atlantic cable, and William Orton, president of Western Union, for free employment of the Atlantic Cable and the telegraph system in the United States for the transmission of astro- nomical data. By February 1873 the Smithsonian had be- gun transmitting information to the Royal Greenwich Ob- servatory for further dissemination to Europe, and through the Associated Press, to astronomers throughout the United States. On the other side of the Atlantic, the European state telegraph companies eventually also agreed to carry the data free of charge. By May 1873 that Henry was able to announce the launching of the system, with European nodes at the major national observatories: Greenwich, Paris, Ber- lin, Vienna, and, a little later, Pulkova. Working out some of the confusion over which of the observatories had what re- porting responsibilities took some time to work out, as did developing a standard lexicon, but by 1883, when Spen- cer Baird, Henry?s successor at the Smithsonian, turned the responsibility for the U.S. node over to Harvard College Observatory, the information exchange was world-wide. Approximately fi fty European observatories were linked to Harvard?s counterpart in Europe, the observatory at Kiel, and connections had also been made with observatories in South America, Australia, and South Africa (Rothenberg et al., 2007, 448; Jones and Boyd, 1971: 197). TRANSITS OF VENUS Transits of Venus, the observation from Earth of the passage of that planet across the face of the sun, are rare astronomical events. Two occur eight years apart, with a gap of over a century between pairs. Because of their application in establishing the astronomical unit, the dis- tance between the earth and the sun, which is the essen- tial yardstick for solar system astronomy, the astronomi- cal community was very eager to take advantage of the opportunities provided by the transits of 1874 and 1882. Ultimately, 13 nations sent out observing expeditions to observe one or both transits. A number of nations estab- lished government commissions to oversee the efforts, in- cluding the United States. Astronomers exchanged copies of their observing protocols and coordinated with each other in selecting observing sites (Dick, 2004; Duerbeck, 2004; Dick, 2003: 243, 265). Planning had begun as early as 1857, with the publi- cation of Astronomer Royal George B. Airy?s suggestions of possible observing sites (Airy, 1857). Among the de- sirable locations, from an astronomical perspective, was Antarctica. For the fi rst time, there was serious discussion of establishing a scientifi c observing site in Antarctica. But the transits occurred in December. Were astronomi- cal observations in Antarctica that time of year practical? Scientists were divided. Airy, using information provided him from Edward Sabine, concluded that ?December is rather early in the season for a visit to this land, but prob- ably not too early, as especially fi rm ice will be quite as good for these observations as dry land? (1857: 216). He called for a reconnaissance ahead of time to test whether it was practical to establish an observing station in the Polar Re- gions. J. E. Davis, a British naval offi cer and Arctic explorer, was even more optimistic, although very realistic as to the diffi culties. He developed a plan in 1869 for observations of the 1882 transit from Antarctica, but noted in his presenta- tion to the Royal Geographical Society (1869), that such observations would have required the observing parties to winter over. There was insuffi cient time to fi nd a safe har- bor and establish the observing station prior to the transit. In the case of the 1882 observers, Davis argued that they should be landed in late 1881 with suffi cient supplies to last two years, even though the plan was to have them picked up in about a year. It was necessary to leave a margin of error. He did warn of the problematic weather conditions, describing the weather as ?either very bad or very delight- ful? (1869: 93). To Davis, it was a gamble worth taking, but it seemed less attractive to astronomers who were going to be making once in a lifetime observations. In contrast to Davis and Airy, Simon Newcomb, the leading American astronomer and a member of the American Transit of Ve- nus Commission, was much more pessimistic. He rejected the idea of astronomical observations from ?the Antarctica continent and the neighboring islands . . . because a party 02_Rothenberg_pg013-000_Poles.in17 1702_Rothenberg_pg013-000_Poles.in17 17 11/17/08 8:36:50 AM11/17/08 8:36:50 AM 18 SMITHSONIAN AT THE POLES / ROTHENBERG can neither be landed nor subsisted there; and if they could, the weather would probably prevent any observations from being taken? (1874: 30). Although observations were not made from the con- tinent of Antarctica, the 1874 transit was observed by parties from Britain, Germany, France, and the United States from stations on islands within the Antarctic Con- vergence, including Kerguelen (Newcomb apparently thought Kerguelen suffi ciently north not to be considered ?a neighboring island?) and Saint-Paul. The 1874 observa- tions were only moderately successful because of weather problems (Bertrand, 1971: 258; Duerbeck, 2004;). But the seeds were planted for a more extensive investigation of the Antarctic. Davis had argued from the beginning that the Antarctic stations should also ?obtain a series of ob- servations in meteorology and other branches? (1869: 93), while the American expedition conducted biological and geological collecting which ?resulted in a signifi cant contribution to the scientifi c knowledge of the Antarctic? (Bertrand, 1971: 255). Although the combination of the uncertainty of the weather and the diffi culties, dangers, and expense of sending parties to Antarctica seemed to have discouraged most further efforts in that direction for the 1882 transit, Germany sent an expedition to South Georgia for the dual purpose of conducting transit of Ve- nus observations and other observations as part of the IPY (Duerbeck, 2004: 14). Later observers have recognized the signifi cance of the transit of Venus expeditions in establishing a prece- dent for later cooperative research in Antarctica. Julian Dowdeswell of the Scott Polar Research Institute has called these observations of 9 December 1872, ?the earliest ex- ample of international coordination in polar science and a clear precursor to the fi rst IPY? (Dowdeswell, 2007). The geographer Kenneth Bertrand also argues that the transit observations belong to the history of Antarctic research, although he skips over the IPY, because it was primarily an Arctic venture, and contends that the international pro- gram for observing the transit was a ?predecessor of the International Geophysical Year? (1971: 255). POLAR STUDIES: SURVIVING THE ELEMENTS AND MORE So if the fi rst IPY was not a path breaking forerunner of later international cooperative research programs, what was its signifi cance? It was ?Polar.? That may seem obvi- ous, but at the time when it was organized, uncertainty hung over Polar research, at least in the United States. The question might be asked: would any effort to gather data in the Polar Regions be a waste of human and scientifi c resources? The United States, and especially the Smithsonian, had supported scientifi c research in the Arctic during the three decades prior to the IPY (Lindsay, 1993; Sherwood, 1965; Fitzhugh, this volume). Some of this could be sol- idly placed under the heading of international coopera- tion, though not at the level exemplifi ed by the fi rst IPY. For example, the Smithsonian had developed strong ties with the British Hudson?s Bay Company, and in a spirit of international cooperation, company employees had col- lected natural history specimens and made meteorological observations. In addition, Francis L. McClintock, a British Polar explorer, had turned his Arctic meteorological ob- servations over to the Smithsonian for reduction (Rothen- berg et al., 2004: 142, 143). The Smithsonian had also arranged for the reduction and publication of the geophysical observations made by two U.S. polar endeavors, the second Elisha Kent Kane expedition (1853? 1855) and the I. I. Hayes expedition (1860? 1861). The apparatus used in the latter expedi- tion were on loan from the Smithsonian (Rothenberg et al., 2004: 142? 144). In addition, the Smithsonian encour- aged natural history research at relatively lower latitudes in Alaska as part of its broader program of supporting the scientifi c exploration of the American West (Fitzhugh, this volume; Goetzmann, 1966). Among the collectors were Robert Kennicott, working in conjunction with the West- ern Union Telegraph Company?s survey of a telegraph route across Alaska, and W. H. Dall, who was Kennicott?s successor and then served with the U.S. Coast Survey (Rothenberg et al., 2007: 128, 397). Both were very closely associated with the Smithsonian and its northern research and collecting program (Fitzhugh, this volume). Polar research was dangerous, as Henry admitted in 1860, requiring ?much personal inconvenience and per- haps risk of life? (Rothenberg et al., 2004: 141). That opinion was no doubt further reinforced by the death of Kennicott in 1866 and the disaster of the U.S. Polaris Ex- pedition, led by Charles Francis Hall, in 1871. This latter expedition has been renown in the history of exploration because of the debate over whether the expedition?s scien- tist/physician murdered the commander (Loomis, 1991). But beyond the human cost, the expedition?s failure tem- porarily dashed the hopes of the American scientifi c com- munity for governmental support for intensive research in the Polar region. Although it has been claimed that Hall, an experienced Arctic explorer, was ?lacking credibility as a man of sci- 02_Rothenberg_pg013-000_Poles.in18 1802_Rothenberg_pg013-000_Poles.in18 18 11/17/08 8:36:51 AM11/17/08 8:36:51 AM COOPERATION AT THE POLES? 19 ence? (Robinson, 2006: 76), he had received the endorse- ment of Joseph Henry during the debate over who would lead the expedition (Rothenberg et al., 2007: 288). There was no question that science was to be a part of the expedi- tion. The legislation which established the expedition, and provided an appropriation of $50,000 for it, ordered that ?the scientifi c operations of the expedition be prescribed in accordance with the advice of the National Academy of Sciences? (United States, 1871: 251). That advice, including the selection of the scientist, Dr. Emil Bessels, a zoologist, came primarily from Henry, as president of the National Academy of Sciences, and his Assistant Secretary at the Smithsonian, Spencer F. Baird, who chaired the Academy committee for the expedition. In his report on the prepa- ration for the scientifi c aspect of the expedition, Henry acknowledged that Hall?s primary mission was ?not of a scientifi c character? and that to have attached ?a full corps of scientifi c observers? to it would have been inappropriate (Rothenberg et al., 2007: 352). Offi cially, he recognized the reality of the politics of exploration and was willing to settle for having Bessels and a few junior observers on the expedi- tion, armed with instructions from some of the leading sci- entists in the United States on collecting data and specimens in astronomy, geophysics, meteorology, natural history, and geology. Unoffi cially, Henry had the expectation that if the Polaris Expedition was successful, Congress could be persuaded to follow up the triumph with an appropriation ?for another expedition of which the observation and in- vestigation of physical phenomena would be the primary object? (Rothenberg et al., 2007: 355). But with the failure of the Polaris Expedition in 1871, the hope of additional Congressional funding was dashed. It would be another de- cade before another U.S. government-sponsored expedition would be sent to the polar region, and it would occur under the auspices of the fi rst IPY in 1881. Participation by the United States in the fi rst IPY in 1881? 1884 was coordinated by U.S. Army Signal Corps, which was experienced in conducting meteorological observations. However, the Smithsonian was in charge of many aspects of the U.S. IPY scholarly program and laid claim to all its resulting ?natural history? collections (Krupnik, this volume). The eastern U.S. mission, on Elles- mere Island, under the command of Lt. Adolphus Greely, was a reasonable success from the perspective of its scien- tifi c observations and data returned. But the expedition was plagued by mutiny, bad luck, and poor judgment, and more than two-thirds of the participants perished. In con- trast, the Alaskan (Point Barrow) mission, commanded by Lt. P. Henry Ray, had an uneventful time, returning valuable scientifi c data and abundant natural history and ethnology collections with little fuss (Burch, this volume; Crowell, this volume; Krupnik, this volume). Furthermore, the little fuss that accompanied Ray?s expedition may be the most important aspect of it and the reason why it was a turning point in the history of American scientifi c ven- tures in the polar regions. For the fi rst time, an expedi- tion ?made survival in the Arctic wastes at 70? below zero look routine to Americans? (Goetzmann, 1986: 428). That survival was at least a reasonable expectation was a neces- sary premise to further scientifi c exploration of the polar regions. CONCLUSION There is an important caveat to this apparent success story of increasing science cooperation. An agreement for international cooperation was not always followed by implementation. As one representative to the unsuccess- ful 1853 Brussels conference noted, when the delegates returned home, ?every one followed his own plan and did what he pleased? (Fleming, 1990: 109). Even after the founding of the International Meteorological Organiza- tion in 1879, confl ict was avoided by the issuing of ?reso- lutions and recommendations that national weather ser- vices could, and often did, ignore? (Edwards, 2004: 827). In addition, William Budd (2001) was correct in iden- tifying the same half century between roughly 1835 and 1885, which I argue was marked by increased efforts at scientifi c cooperation, as also a half century of intense international rivalry. And that rivalry is the fl ip side of the history of scientifi c cooperation. Prestige and glory were strong motivations for participating in collabora- tive ventures. National scientifi c communities could, and frequently did, point to the activities of international ri- vals to encourage their governments to provide fi nancial support for research. Participation in certain international endeavors, such as the Magnetic Crusade or the transit of Venus observations, the argument went, was absolutely necessary if a nation was to maintain status within the international community. Scientists would use the activi- ties of other governments to shame their own to action. As Cawood noted, the willingness of the Norwegian Parlia- ment to fund a geomagnetic expedition in 1828, while at the same time denying the funds for the erection of a royal residence, ?became an almost obligatory precedent to be quoted in all British pleas for the government backing of terrestrial magnetism? (1979: 506). Cawood warned, moreover, that there was ?a very narrow dividing line be- tween international cooperation and international rivalry? 02_Rothenberg_pg013-000_Poles.in19 1902_Rothenberg_pg013-000_Poles.in19 19 11/17/08 8:36:51 AM11/17/08 8:36:51 AM 20 SMITHSONIAN AT THE POLES / ROTHENBERG (1979: 518). When international relations soured, or na- tionalistic emotions increased, the presence of government funding and the recognition of the domestic political value of scientifi c success could possibly result in national fac- tors overwhelming the cooperative, international aspects of research. International cooperation in meteorology suf- fered from the unwillingness of government-funded scien- tists to turn their backs on the immense investment they had made in their own systems and accept a foreign sys- tem. It remains to see, when historians look back, whether rivalry or cooperation will be the dominant theme for the latest International Polar Year in 2007? 2009. LITERATURE CITED Airy, George B. 1857. On the Means which Will Be Available for Cor- recting the Measure of the Sun?s Distance, in the Next Twenty-Five Years. Monthly Notices of the Royal Astronomical Society, 17: 208? 221 Anderson, Katharine. 2005. Predicting the Weather: Victorians and the Science of Meteorology. Chicago: Chicago University Press. Barr, William. 1983. Geographical Aspects of the First International Po- lar Year, 1882? 1883. Annals of the Association of American Geog- raphers, 73: 463? 484. Bertrand, Kenneth J. 1971. Americans in Antarctica, 1775? 1948. New York: American Geographical Society. Budd, W. F. 2001. ?The Scientifi c Imperative for Antarctic Research.? In The Antarctic: Past, Present and Future, ed. J. Jabour-Green and M. Haward, pp. 41? 59. Antarctic CRC Research Report 28. Burch, Ernest S., Jr. 2009. ?Smithsonian Contributions to Alaskan Eth- nography: The IPY Expedition to Barrow, 1881? 1883.? In Smith- sonian at the Poles: Contributions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 89? 98. Washington, D.C.: Smithsonian Institution Scholarly Press. Cawood, John. 1977. Terrestrial Magnetism and the Development of In- ternational Collaboration in the Early Nineteenth Century. Annals of Science, 34:551? 587. ???. 1979. The Magnetic Crusade: Science and Politics in Early Vic- torian Britain. Isis, 70:493? 518. Crowell, Aron L. 2009. ?The Art of I?upiaq Whaling: Elders? Interpre- tations of International Polar Year Ethnological Collections.? In Smithsonian at the Poles: Contributions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 99? 114. Washington, D.C.: Smithsonian Institution Scholarly Press. Davis, J. E. 1869. On Antarctic Discovery and Its Connection with the Transit of Venus. Journal of the Royal Geographical Society, 39: 91? 95. Dick, Steven J. 2003. Sky and Ocean Joined: The U.S. Naval Observa- tory, 1830? 2000. Cambridge, U.K.: Cambridge University Press. ???. 2004. The American Transit of Venus Expeditions of 1874 and 1882. In Proceedings of IAU Colloquium No. 196, pp. 100? 110. Cambridge, U.K.: Cambridge University Press. Dowdeswell, Julian. U.K.? Scott Polar Research Institute, International Polar Year Outreach Project. http://www.spkp.se/ipy/ipyou03.htm (accessed 7 September 2007). Duerbeck, Hilmar W. 2004. The German Transit of Venus Expeditions of 1874 and 1882: Organizations, Methods, Stations, Results. Journal of Astronomical History and Heritage, 7: 8? 17. Edwards, Paul N. 2004. ?A Vast Machine?: Standards as Social Technol- ogy. Science, 304: 827. Fitzhugh, William W. 2009. ??Of No Ordinary Importance?: Revers- ing Polarities in Smithsonian Arctic Studies.? In Smithsonian at the Poles: Contributions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 61? 78. Washington, D.C.: Smithsonian Institution Scholarly Press. Fleming, James Rodger. 1990. Meteorology in America, 1800? 1870. Bal- timore: Johns Hopkins University Press. Goetzmann, William H. 1966. Exploration and Empire: The Explorer and the Scientist in the Winning of the American West. New York: Alfred A. Knopf. ???. 1986. New Lands, New Men: America and the Second Great Age of Discovery. New York: Viking. Hawes, Joseph M. 1966. The Signal Corps and Its Weather Service, 1870? 1890. Military Affairs, 30: 68? 76. Jones, Bessie Zaban, and Lyle Gifford Boyd. 1971. The Harvard Col- lege Observatory: The First Four Directorships. Cambridge, Mass.: Harvard University Press. Krupnik, Igor. 2009. ??The Way We See It Coming?: Building the Legacy of Indigenous Observations in IPY 2007? 2008.? In Smithsonian at the Poles: Contributions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 129? 142. Washington, D.C.: Smithsonian Institution Scholarly Press. Lindsay, Debra J. 1993. Science in the Subarctic: Trappers, Traders, and the Smithsonian Institution. Washington, D.C.: Smithsonian Insti- tution Press. Loomis, Chauncy C. 1991. Weird and Tragic Shores: The Story of Charles Francis Hall, Explorer. Lincoln: University of Nebraska Press. Luedecke, Cornelia. 2004. The First International Polar Year (1882? 83): A Big Science Experiment with Small Science Equipment. History of Meteorology, 1: 55? 64. [Newcomb, Simon]. 1874. The Coming Transit of Venus. Harper?s New Monthly Magazine, 50: 25? 35. Reingold, Nathan, Arthur P. Molella, Marc Rothenberg, Kathleen Waldenfels, and Joel N. Bodansky, eds. 1979. The Papers of Joseph Henry. Volume 3. Washington, D.C.: Smithsonian Institution Press. Rhees, William J. 1859. An Account of the Smithsonian Institution, Its Founder, Building, Operations, Etc., Prepared from the Reports of Prof. Henry to the Regents, and Other Authentic Sources. Washing- ton, D.C.: Thomas McGill. Robinson, Michael F. 2006. The Coldest Crucible: Arctic Exploration and American Culture. Chicago: University of Chicago Press. Rothenberg, Marc. 1999. Peters, Christian Henry. American National Biography, 17: 391? 392. Rothenberg, Marc, Kathleen W. Dorman, Frank R. Millikan, Deborah Y. Jeffries, and Sarah Shoenfeld, eds. 2004. The Papers of Joseph Henry. Volume 10. Sagamore Beach, Mass.: Science History Pub- lications. ???, eds. 2007. The Papers of Joseph Henry. Volume 11. Sagamore Beach, Mass.: Science History Publications. Sabine, Edward. 1866. Note on a Correspondence between Her Majesty?s Government and the President and Council of the Royal Society Regarding Meteorological Observations to Be Made by Land and Sea. Proceedings of the Royal Society of London, 15: 29? 38. Sherwood, Morgan. 1965. Exploration of Alaska, 1865? 1900. New Haven, Conn.: Yale University Press. Smithsonian Institution. 1848. Annual Report of the Board of Regents of the Smithsonian Institution for 1847. Washington, D.C.: Tippen and Steeper. ???. 1872. Annual Report of the Board of Regents of the Smithson- ian Institution for 1868. Washington, D.C.: Government Printing Offi ce. 02_Rothenberg_pg013-000_Poles.in20 2002_Rothenberg_pg013-000_Poles.in20 20 11/17/08 8:36:51 AM11/17/08 8:36:51 AM COOPERATION AT THE POLES? 21 Taylor, C. J. 1981. First International Polar Year, 1982? 83. Arctic, 34: 370? 376. United States. 1871. The Statutes at Large and Proclamations of the United States of America, from December 1869 to March 1871. Boston: Little, Brown, and Company. United States, Commission to the Paris Universal Exposition. 1880. Re- ports of the United States Commissioners to the Paris Universal Exposition, 1878. Volume 1. Washington, D.C.: Government Print- ing Offi ce. Weiss, Edith Brown. 1975. International Responses to Weather Modifi - cation. International Organization, 29: 805? 826. Williams, Francis L. 1963. Matthew Fontaine Maury, Scientist of the Sea. New Brunswick, N.J.: Rutgers University Press. 02_Rothenberg_pg013-000_Poles.in21 2102_Rothenberg_pg013-000_Poles.in21 21 11/17/08 8:36:52 AM11/17/08 8:36:52 AM 02_Rothenberg_pg013-000_Poles.in22 2202_Rothenberg_pg013-000_Poles.in22 22 11/17/08 8:36:52 AM11/17/08 8:36:52 AM ABSTRACT. By the post? World War II era, the U.S. federal government?s role in science had expanded considerably. New institutions, such as the Offi ce of Naval Research and the National Science Foundation, were established to fund basic science. Technologi- cal breakthroughs that had provided the instruments of war were recognized as having important economic, civilian applications. Understanding the earth?s environment, in- cluding the extreme polar regions, the upper atmosphere, and the ocean depths, was rec- ognized as key to enhancing a nation?s communications, transportation, and commerce. The IGY developed in part from such national interests, but became a huge international undertaking. The process of international negotiations leading up to and during the IGY set a precedent for organizing cooperative scientifi c undertakings and enshrined norms and practices for sharing data and resources. Further, the IGY demonstrated the impor- tance of communicating results across political, disciplinary, and societal boundaries. Fifty years later, the organizers of the International Polar Year embraced these values. INTRODUCTION Legacies of the Third International Polar Year, more commonly known as the International Geophysical Year (IGY) of 1957? 1958, include the launch of the fi rst artifi cial Earth-orbiting satellites, the negotiation of the Antarctic Treaty, the establishment of the World Data Center system, the discovery of the Van Allen belts, and the long-term measurements of atmospheric carbon dioxide and glacial dynamics (Sullivan 1961; Korsmo 2007b). While the outcomes of the IGY are well known, the social and political processes that led to the IGY are less studied. One of the best sources from a policy perspective is a small pamphlet produced by the Congressional Research Service (Bullis 1973). An- other source is the U.S. National Academy of Sciences voluminous IGY Archive in Washington, D.C., in addition to other archives containing the papers of IGY scientists and government sponsors. Some IGY participants have contributed their recollections in oral histories (e.g., Van Allen, 1998; see Acknowledge- ments), an excellent source of information on what it was like to be a researcher or an administrator before, during, and after the IGY. This chapter draws on archival collections, biographies, oral histories, and secondary sources to ask the question, What lessons can IGY teach us about Fae L. Korsmo, National Science Foundation, Offi ce of the Director, Arlington, VA 22230, USA (fkorsmo@nsf.gov). Accepted 29 May 2008. The Policy Process and the International Geophysical Year, 1957? 1958 Fae L. Korsmo 03_Korsmo_pg023-034_Poles.indd 2303_Korsmo_pg023-034_Poles.indd 23 11/17/08 8:38:17 AM11/17/08 8:38:17 AM 24 SMITHSONIAN AT THE POLES / KORSMO organizing and carrying out a large-scale, international, and multidisciplinary science program? The IGY helped to establish a precedent for the post? World War II conduct of international science, particularly as organized cam- paigns of international years or decades. In addition, the U.S. organizers of the IGY within the National Academy of Sciences left a well-organized archive of documentation, including a set of full transcripts from executive committee meetings. It is as if they wanted to be studied, since they purposely recorded their daily conversations, arguments, and decisions. As a political scientist, I fi nd this record irre- sistible. When it comes to setting science agendas, building coalitions to advocate for programs, and making sure that commitments for funding and other support are made and kept, it is seldom one has access to so complete a record of events. Even as we fi nd ourselves in the midst of the [Fourth] International Polar Year 2007? 2008, it is not too late to learn from the IGY. Policy can be thought of as a set of processes, includ- ing the setting of the agenda (the list of problems or sub- jects to which the decision-makers are paying serious at- tention at any given time); the specifi cation of alternatives from which a choice is to be made; an authoritative choice among the alternatives, and the decision itself (Kingdon, 1995). In the process of setting the agenda or advocating for alternatives, those motivated by the outcomes form coalitions and engage in coordinated behavior based on common beliefs or motivations (Sabatier and Jenkins- Smith, 1999). Policy entrepreneurs? advocates willing to invest their resources to promote an alternative in return for anticipated future gain? look for opportunities to link solutions to problems, using what they know about the relevant coalitions and decision-makers. They are the clas- sic integrators, creating connections among people, prob- lems, alternatives, and decisions. Once a decision is made, then the question becomes, Who will implement the deci- sion? There must be a credible commitment among insti- tutions (and by ?institutions,? I mean the rules and norms that groups of people live by, which may or may not be embodied in organizations) to match subsequent actions with authoritative choices (North and Weingast, 1989). In the follow-through, the rules of the game and lines of accountability become crucial. The IGY, on the one hand a daring and audacious plan to make the entire earth? surface, oceans, and at- mosphere? the topic of concerted studies, and on the other hand a series of small, incremental decisions taken independently by numerous small groups, can be under- stood as a set of policy processes. A retrospective analysis of what worked enabled us to compare these IGY lessons with the science planning we are doing now. Was the IGY simply a product of its time or are there enduring legacies in terms of how we construct and conduct coordinated research programs? In terms of process, this chapter pro- vides some of the highlights. First, however, it is helpful to examine the political context of the IGY in comparison to the other Polar Years. THE POLITICAL CONTEXT All of the Polar Years occurred in periods of relative peace and political stability, when international organiza- tions had emerged in a new or reconstituted fashion (e.g., from periods of inactivity during wartime or economic de- pression) and when powerful nations were not distracted by the drain of soldiers and armaments. The idea for a First Polar Year or coordinated polar studies, for example, emerged within the Austro-Hungarian Empire in the early 1870s but only gained traction with other major powers after the 1878 Congress of Berlin had settled the Balkan War (Baker, 1982). The idea for a Second Polar Year was raised in 1927 and the studies took place in 1932? 1933, fi fty years after the First Polar Year (Baker, 1982; Nicolet, 1984). The Great Depression, followed by the outbreak of World War II and the untimely death of the International Polar Year Commission?s President D. LaCour in 1942 re- sulted in a lengthy delay in publishing the results from the Second Polar Year. By the end of World War II, science and technology, as well as politics, had changed. The main fac- tor that separates the IGY from the fi rst two Polar Years was the existence of nuclear weapons and the beginnings of the Cold War between East and West. Understanding, detection, and development of nuclear weapons became a driving force for investment in the physical sciences fol- lowing World War II. The Arctic, as geographer Paul Siple pointed out in 1948, afforded a straight line of attack to the Soviet Union (Siple, 1948). The United States began to con- duct top-secret overfl ight missions to determine whether the Soviet forces were staging long-range bombers in the fro- zen north (Hall, 1997). The artifi cial Earth-orbiting satel- lites that emerged from the IGY were useful in studying the upper atmosphere but also provided a means of checking on enemy activities (Day, 2000). Basic science and military objectives both were motivations. The debate among histo- rians has centered on how inextricably linked the motiva- tions were for participating scientists and institutions. As in the fi rst two Polar Years, international scientifi c organiza- tions, such as the International Meteorological Congress in the 1870s and the International Council of Scientifi c Unions 03_Korsmo_pg023-034_Poles.indd 2403_Korsmo_pg023-034_Poles.indd 24 11/17/08 8:38:17 AM11/17/08 8:38:17 AM POLICY PROCESS AND THE INTERNATIONAL GEOPHYSICAL YEAR 25 in the 1930s, played a key role in providing stable, reliable fora in which to build coalitions and collaborations. How- ever, individuals? our policy entrepreneurs? also were re- quired to initiate and carry out ambitious programs. AGENDA SETTING: FIRST, SET A DATE It sounds easy to set a date but even this requires some thought. How many declared international years of this and decades of that pass unnoticed? There has to be a rea- son for the date, preferably a reason tied to the proposed activity, an agenda that establishes the urgency of action within a specifi c time frame. What is the scientifi c justi- fi cation for an ?international year?? Unless the urgency appeals to a community of scientists beyond the initially small group of advocates, it will be diffi cult to affi x a con- vincing time period for intense activity. Setting a date, then, requires the ability to persuade others and build coalitions of advocates and future per- formers? those who will carry out the activity. Building the coalitions requires access to other potential support- ers who can help to make the case. Access comes through networks of contacts, including committees, boards, and other fora. It helps to start with venues that allow for the ex- change of ideas. Most sources place the beginning of the IGY at the home of Dr. James Van Allen, in Silver Spring, Maryland in the spring of 1950 (Sullivan, 1961; Van Al- len, 1997 and 1998). Trained as a nuclear physicist, Van Allen (1914? 2006) was well known for the development of the proximity fuze during World War II. He became involved in the use of rockets to study the upper atmo- sphere immediately following the war, instrumenting captured and refurbished German V-2 rockets to study cosmic radiation, the ionosphere, and geomagnetism. At the time of Chapman?s visit, Van Allen headed up a high- altitude research group at Johns Hopkins University, Ap- plied Physics Lab. Shortly thereafter, Van Allen would go to the University of Iowa, where he would spend most of his professional career as a professor of physics (American Institute of Physics, 2000). While in Maryland, he and his wife Abigail hosted a dinner on 5 April 1950 for British geophysicist Sydney Chapman (1888? 1970). Chapman, a theoretical physicist interested in the earth?s magnetic phenomena, had participated in the Second Polar Year of 1932? 1933. He was well known for his work on magnetic storms, and would come to spend a great deal of time in the United States, at the University of Alaska Fairbanks, the High Altitude Observatory in Boulder, Colorado, and University of Michigan (Good, 2000). He was to start his lengthy U.S. sojourn at Caltech. In April 1950, Chapman was in the United States on his way to join a Caltech study on the upper atmosphere. Van Allen described the gath- ering as ?one of the most felicitous and inspiring? that he had ever experienced. Also present at the dinner was Lloyd Berkner, a former radio engineer who had been on Admiral Byrd?s 1928? 1930 Antarctic expedition. Berkner had both science and policy in his background. According to Van Allen, The dinner conversation ranged widely over geophysics and especially geomagnetism and ionospheric physics. Following dinner, as we were all sipping brandy in the living room, Berkner turned to Chapman and said, ?Sydney, don?t you think that it is about time for another international polar year?? Chapman im- mediately embraced the suggestion, remarking that he had been thinking along the same lines himself. (Van Allen, 1998:5) Chapman also observed that the years 1957? 1958 would be a time of maximum solar activity, so the time frame for the Third Polar Year was settled. The proper- ties of the upper atmosphere, including the relationships among magnetic storms, cosmic rays, and solar activity in- trigued scientists. The military, in particular, was interested in very-high-frequency scatter technology for reliable low- capacity communication that could avoid the disruptions caused by solar emissions, magnetic storms, and auroras. The perturbations emanated from high latitudes. Choosing a year of maximum solar activity for a new international polar venture made scientifi c sense for atmospheric physi- cists interested in understanding more about these high- latitude phenomena. Furthermore, as Chapman explained later (1960, 313) and may well have noted at the Van Allen residence, technological improvements in instrumentation and rocketry had now enabled scientists to probe much deeper into the atmosphere. The time was ripe. Was the Berkner-Chapman exchange rehearsed? Perhaps. But the main point is that there were many opportunities for lead- ing scientists to get together and persuade one another of the need for an intense research campaign (Kevles, 1990). One of the main venues in the United States after World War II was the Joint Research and Development Board, re- named the U.S. Research and Development Board in 1947. Led by Vannevar Bush, the Research and Development Board combined civilian researchers and military person- nel in determining and coordinating research priorities for the Department of Defense. Bush was President of the Carnegie Institution of Washington and had led the Joint Committee on New Weapons and Equipment during the 03_Korsmo_pg023-034_Poles.indd 2503_Korsmo_pg023-034_Poles.indd 25 11/17/08 8:38:18 AM11/17/08 8:38:18 AM 26 SMITHSONIAN AT THE POLES / KORSMO war. Widely respected today as the author of Science? The Endless Frontier (Bush, 1945), a book that stressed the im- portance of basic research in the United States, Bush repre- sented the transition from wartime science? applications focused on immediate problems of winning the war? to peacetime science? research into fundamental questions for the sake of discovery. The Research and Development Board?s committees and their subcommittees and panels took up scientifi c problems that the military identifi ed? or civilian scientists identifi ed for them. They covered the physical, medical, bi- ological, and geophysical sciences (U.S. National Archives and Records Administration, n.d.). Vannevar Bush tapped Lloyd Berkner (1905? 1967) to run the Research and De- velopment Board as its executive secretary. Berkner, pro- fi led in Allan Needell?s excellent biography (2000), had a rich experience in ionospheric research, government con- sulting, and national security. He had been to Antarctica in 1928? 1930. He was not afraid to speak out publicly on science policy. He was perfect example of a policy entre- preneur: breaking down boundaries between government agencies and between government and other sectors of society, exploiting every opportunity for bringing solu- tions? in this case technological breakthroughs and sci- entifi c programs? to problems. In the world of agenda setting, solutions, carried in the pockets of entrepreneurs, go searching for problems. While we are left with the im- pression that Berkner fi rst introduced the idea of a Third International Polar Year at the Van Allen home in April 1950, undoubtedly he had broached the topic before, per- haps on the Research and Development Board, using his previous Antarctic experience and his knowledge of post- war developments in international science policy. Berkner was not the only science entrepreneur in the postwar science world. One can also think of Swedish mete- orologist Carl Gustav Rossby (1898? 1957), who called the attention of the U.S. military to the climate warming occur- ring in the 1920s through the 1940s (e.g. Rossby 1947). His fellow Swede, glaciologist Hans W. Ahlmann (1889? 1974), studied the properties of glaciers as indicators of climate variation. Rossby urged the U.S. military to take advan- tage of Ahlmann?s knowledge as the military searched for expertise on high-latitude operations. Because the Arctic represented the shortest distance between the United States and Soviet territory, Rossby and Ahlmann knew they had the attention of the U.S. military planners. Using solid ice for planes and other military transport was fi ne as long as the ice was not melting; a warming trend would require a change in strategy. In a way, it was Ahlmann who helped to set the science agenda for the IGY?s approach to glaciol- ogy. Using mountain glaciers, Ahlmann measured accumu- lation, ablation, and regime (the grand total of a glacier?s entire accumulation and net ablation) and recommended simultaneous measurements in different climates (Kirwan et al., 1949). In 1946, he called for exact measurements to be made on the ice sheets: temperatures at different depths, seismic methods to compute thickness, and detailed obser- vations of stratifi cation of annual layers. The only way to do this systematic comparison in different parts of the world was through inter national cooperation (Ahlmann, 1946). Ahlmann?s frequent calls for this style of comparative work eventually led to the Norwegian- British-Swedish Antarc- tic Expedition of 1949? 1952, the fi rst scientifi c traverse in the Antarctic interior. This traverse served as a model for the IGY expeditions to Antarctica (Bentley, 1964). Of course, the U.S. had political reasons for a scientifi c pres- ence in Antarctica; the existence of competing territorial claims and concerns about possible Soviet claims ensured the Antarctic would have a place on the IGY agenda (U.S. National Security Council, 1957). How did the Third Polar Year, originally envisioned as a high-latitude, upper-atmosphere research campaign, become the International Geophysical Year? In the process of enlisting support among the international scientifi c soci- eties, Chapman and Berkner found a strong preference for a global program encompassing additional geographical regions and physical science disciplines. To attain wide- spread support, Chapman and Berkner skillfully embraced a much broader geophysical agenda. The international science scene after World War II in- cluded a reconstituted International Council of Scientifi c Unions (ICSU), a body where membership was both by nation-state and by international scientifi c union, e.g., the International Union of Geodesy and Geophysics (IUGG). The national member might be a national academy or a government agency with research responsibilities. Berkner and Chapman fi rst presented the idea for the Third In- ternational Polar Year to the constituent scientifi c unions that made up a ?Mixed Commission on the Ionosphere? under ICSU (Beynon, 1975:53). These unions included the IUGG, International Astronomy Union (IAU), Inter- national Union of Pure and Applied Physics (IUPAP), and International Union of Radio Science (URSI) (Beynon, 1975:53). The unions, in turn, presented the proposal to the ICSU General Assembly, and ICSU, in turn, invited the World Meteorological Organization (WMO) to par- ticipate as well as the national organizations adhering to ICSU. (Note that this pattern of approaching interna- 03_Korsmo_pg023-034_Poles.indd 2603_Korsmo_pg023-034_Poles.indd 26 11/17/08 8:38:18 AM11/17/08 8:38:18 AM POLICY PROCESS AND THE INTERNATIONAL GEOPHYSICAL YEAR 27 tional scientifi c organizations fi rst ICSU, then the WMO, was also used to prepare for the International Polar Year 2007? 2008). By 1953, there were 26 countries signed up to participate in what came to be known as the Inter- national Geophysical Year 1957? 1958. The disciplines in- cluded practically all the earth, atmosphere, and oceanic sciences, covering many parts of the globe beyond the polar regions (Nicolet, 1984). The price of coalition build- ing beyond the minimum one needs to win an objective is that the agenda? the set of interesting science questions and topics? becomes expansive and unwieldy. Political scientists have theorized that a minimum winning coalition? the number of political parties, repre- sentatives, or individuals needed to win a particular com- petition such as an election? will prevail in democratic politics given a zero-sum situation of clear winners and losers (Riker, 1962). Surplus coalition members may bring instability and demands that cannot be met. While coali- tion theory has been a lively topic of debate and study in the political science literature (e.g., Cusack et al., 2007), I believe it is a useful concept outside the realm of electoral politics to analyze the building of science coalitions, which are not necessarily zero-sum endeavors. Did Berkner, Chapman, and their allies know that they needed to build a large umbrella? Their intentions are not explicit, but they may have felt that they needed as much support as possible from the international scientifi c orga- nizations in order to press their case at home for an IGY. To prevent the science agenda from becoming too diffuse, barriers to entry have to be established. By 1954, the inter- national IGY organizing committee (set up by ICSU in 1952 and known as CSAGI after its French name, Comit? Sp?cial de l?Ann?e G?ophysique Internationale) established criteria for IGY proposals. Priority would be given to projects with at least one of the following characteristics: 1. Problems requiring concurrent synoptic observations at many points involving cooperative observations by many nations. 2. Problems in geophysical sciences whose solutions would be aided by the availability of synoptic or other concentrated work during the IGY. 3. Observations of all major geophysical phenomena in relatively inaccessible regions of the Earth that can be occupied during the IGY because of the extraordinary effort during that interval (the Arctic and Antarctic). 4. Epochal observations of slowly varying terrestrial phe- nomena (International Council of Scientifi c Unions, 1959). 1 These were not arbitrary or unreasonable criteria, since they still permitted a variety of disciplines and con- formed in the main with the justifi cation for a coordi- nated program confi ned to an 18-month time period, from July 1957 until December1958. Each discipline fi t- ting the criteria had a reporter, whose responsibilities in- cluded working with the appropriate scientifi c union to organize the program for that discipline. The program for each discipline was fi rst outlined by an IGY Committee created by the appropriate scientifi c union or by some other ICSU body. Detailed coordination of the program, such as the issuance of instruction manuals for the taking of measurements, was the responsibility of the reporter. The overall direction was the responsibility of the CSAGI Bureau (Bullis, 1973; Nicolet, 1984; see Appendix 1). The CSAGI Bureau members, reporters, and members of the CSAGI General Assembly all served based on their scientifi c fi eld and their professional standing in the ICSU unions rather than based on nationality. Representation on the basis of science rather than nationality enabled CSAGI and the committees to focus on the nature of the work to be done. This model set a precedent in subsequent ICSU and joint ICSU/WMO scientifi c campaigns, most recently in IPY 2007? 2008. A separate Advisory Council, composed of one delegate from each national IGY committee, assisted with practical arrangements such as fi nances, regional meetings, access to foreign territory and facilities, bilateral exchanges, and the collection and storage of data. The national com- mittees, through the Advisory Council, were responsible for implementation (Bullis, 1973). The use of national committees for the IGY ensured that government agencies? both as sources of funding and as authorities? would be involved in science plan- ning from the beginning. Decisions about what was to be funded were made at the national level, so that the IGY amounted to a loosely coordinated set of parallel (or si- multaneously run) national programs. The structure of parallel science committees and the advisory council at the international level enabled realistic commitments to be made on the spot. The U.S. National Academy of Sciences assembled the IGY national committee of 19 members (see Appendix 2; Atwood 1952) in early 1953 and it included govern- ment scientists, operational agencies such as the National Weather Bureau, funding agencies, and the military agen- cies that would provide personnel and logistics. The U.S. national committee formed special technical panels to plan the science and evaluate proposals sent to the National Science Foundation. Including the operational agencies 03_Korsmo_pg023-034_Poles.indd 2703_Korsmo_pg023-034_Poles.indd 27 11/17/08 8:38:18 AM11/17/08 8:38:18 AM 28 SMITHSONIAN AT THE POLES / KORSMO ensured that whatever academic scientists had in mind could be compared to what was possible on the ground. The National Science Foundation (NSF), established with a very small budget in 1950, was charged to be the offi cial IGY funding agency in the United States, the one that would carry forward and coordinate the IGY budget for the U.S. government. The director of the NSF, Alan Waterman, enthusiastically supported and lobbied for the IGY, no doubt seeing an opportunity for the small agency to gain visibility and resources (Korsmo and Sfraga, 2003). Internationally, UNESCO helped out fi nancially (Bullis, 1973). As an independent, nonmilitary agency, the NSF offered a credible funding source both here in the United States and internationally, in an era where military fund- ing? accompanied by classifi cation and secrecy? was the prime source of most geophysical research. The IGY was to be civilian in character, with the scientifi c results shared in the open literature. Nevertheless, the logistics for fi eld- work and of course the satellite program would be shoul- dered by the military agencies. What about political and monetary support for the IGY? Where was the money coming from? That is where solutions go searching for problems. In order to persuade nonspecialists to fund the large-scale nonmilitary science program, the U.S. National Committee for the IGY had to move beyond agenda-setting and coalition building with scientists and executive branch agencies to approach the U.S. Congress and respond to congressional concerns about the public value of science. LINK SOLUTIONS TO PROBLEMS: THE STATE OF SCIENCE EDUCATION IN THE UNITED STATES The National Academy of Sciences? IGY Archive pro- vides evidence that the U.S. National Committee for the IGY reached out to many audiences and answered the hundreds of inquiries received from teachers, students, media, and members of the public. The archive contains, for example, a copy of a letter to an elementary school stu- dent who had written of his ambition to become a ?space man.? Hugh Odishaw (1916? 1984), another entrepre- neurial character and, as the Executive Director to the U.S. National Committee, the architect of the Committee?s in- formation strategy, replied with a two-page letter: I was happy to receive your well-written letter, and to learn of your interest in becoming a ?Space Man.? While you are in elementary school, you won?t have to choose your own subjects of course. Later on, when you to go high school and college, you will want to take as many Mathematics and Science courses as possible, together with courses in English, History, and other subjects which will help to make you a well-rounded person, as well as a possible ?Space Man.? The letter goes on to recommend reading all the ma- terials assigned by the student?s teacher in addition to other books such as Ronald Fraser?s Once Round the Sun (H. Odishaw, letter to J. Bunch, 15 April 1958, in Chron. fi le, IGY Offi ce of Information, National Academy of Sci- ences IGY Collection, Washington, D.C.). Odishaw?s response to the young student is typical of the care with which the U.S. National Committee?s Offi ce of Information answered the mail. During the IGY, about a dozen people worked in this Offi ce, maintaining close contact with media, schools, government agencies, Con- gress, private industry, and professional societies. Well before the IGY began in 1957, the U.S. National Committee for the IGY thought about providing educa- tion materials and general information about the excit- ing ?experiments in concert? that were about to begin. Broadly accessible information about the IGY was nec- essary background for the National Science Foundation?s IGY budget request, but also there was a sense of urgency regarding the state of science education in the country. In its 30 November 1956 issue, the magazine U.S. News and World Report focused on education and science, noting the disparity between the Soviet Union and the United States in the size of the workforce engaged in technical and engineering jobs. One of the articles in this issue, by chemist and businessman Arnold Beckman (who de- veloped the fi rst commercially successful pH meter), lev- eled the all-too-familiar complaints about the American school system: it has failed to anticipate and prepare for the increasing need for more scientists and engineers; its science teachers are not competent to teach science; and its teacher certifi cation requirements pay more attention to how to teach rather than mastery of the subject matter (Beckman, 1956). Beckman was not the only one calling for improvements in science and mathematics education across the United States. The U.S. Congress raised the same concerns to the National Science Foundation and the IGY Committee. As recounted to Hugh Odishaw by S. Paul Kramer, a consul- tant to Odishaw who attended the congressional hearings, Senator Everett Dirksen (R? Illinois) urged the IGY scien- tists to involve the high schools and colleges in real time: ?I would not like to see available information embalmed until the year and a half is over,? said Dirksen. ?I would like to 03_Korsmo_pg023-034_Poles.indd 2803_Korsmo_pg023-034_Poles.indd 28 11/17/08 8:38:19 AM11/17/08 8:38:19 AM POLICY PROCESS AND THE INTERNATIONAL GEOPHYSICAL YEAR 29 see it move out where it will do good.? Senator Warren Magnuson (D? Washington) reminded the National Science Foundation that education and public outreach were well within the scope of the Foundation?s mandate: ?I know from having authored the bill. The real reason for approv- ing it was this sort of thing? (S. P. Kramer, correspondence to H. Odishaw, 6 March 1957, in Chron. fi le, IGY Offi ce of Information, National Academy of Sciences IGY Collec- tion, Washington, D.C.). In response, the IGY organizers worked with publishers, universities specializing in teacher training, and organizations such as the Science Service to develop educational pamphlets, teacher guides, posters, and classroom activities. An example of one of IGY posters on the Oceans that also appeared in a pamphlet is reproduced in Figure 1. The IGY organizers also produced a fi lm se- ries, Planet Earth, released in 1960, for use in schools and for educational television (U.S. National Committee for the IGY, 1960; Korsmo, 2004). There was little if any dissent on the need for an edu- cation and information campaign; the organizers believed it was the right thing to do. The arguments that emerged from the archival records were about the best ways of do- ing it (Korsmo, 2004). FULFILLING THE COMMITMENTS: DATA, PUBLICATION, AND RESULTS The justifi cation for having a Third International Polar Year, which became the International Geophysical Year, only twenty-fi ve years after the Second Polar Year of 1932? 1933 rested in part on the many advances made in techniques of geophysical observation, including radio communications and aviation (Chapman, 1960). If new techniques of analysis, including advances in instrumenta- tion and processing, seem adequate to push for large-scale data collection, then how does one go about it on a world- wide scale? Who takes responsibility to collect, process, and share the data in useable formats? This was the question faced by the designers of the In- ternational Geophysical Year. The U.S. National Commit- tee for the IGY met for the fi rst time on 27 March 1953 in Washington, D.C. In Berkner?s absence on the fi rst day, the participants expressed their doubts. There were at least two problems: fi rst, the question of whether the Soviet Union would participate at all, and second, the problem of secrecy and classifi cation of geophysical data that existed in the United States (Gerson, 1953; Needell, 2000). How could you have a worldwide program when the Soviet Union and its allies were not involved? With Stalin?s death earlier that month, on 6 March 1953, the question hung in the air: Would the Soviet Union open up? At the time, the Soviet Union belonged to the World Meteoro- logical Organization and the International Astronomical Union, but not to ICSU or other ICSU member unions. There was hardly any data exchange between the United States and its allies and the Soviet Union except for rou- tine weather observations. On the other hand, the United States also classifi ed much of its data; the entire polar pro- gram funded by the military, for example. How could the United States expect the Soviet Union to supply data when we withheld ours? All high-latitude ionospheric data? the original focus of the Third Polar Year as proposed by Berkner and Chapman back in 1950? were considered classifi ed in the United States. The U.S. National Committee for the IGY was not the only committee to raise the problem of classifi cation and secrecy. In May 1953, the U.S. Research and Development Board?s Geophysics Committee also raised the issue to the army, navy, and air force (U.S. Research and Development Board, 1953). The Geophysics Committee made a distinc- tion between basic data? ?the elementary building blocks of scientifi c progress in the earth sciences?? and the end products, such as reports that might be used for military or national security purposes. Free access to the data, insisted the Committee, was necessary for scientifi c progress. The designers of the IGY deliberately decided in favor of a science program with free and open data exchange. Indeed, they saw IGY as a means to loosen up the secrecy classifi cations in their own country in addition to encour- aging better data fl ow from other nations. By the fall of 1954, it became clear that the Soviet Union would indeed participate in the IGY, so Berkner instructed the U.S. National Committee to prepare a description of all the data that the United States was prepared to gather and exchange. The idea was to get standardized instru- ments to record multiple observations taken at frequent intervals in many parts of the world. These observations would be recorded, analyzed, synthesized, and preserved in usable formats for further study. As Berkner told the U.S. National Committee in 1953, ?Let our measurements be designed so that repeats during the 4th [Geophysical Year] will be valuable? (Gerson, 1953). The idea of the world data centers also came up early in the IGY planning process (Chapman, 1955). The United States volunteered to host one, which became a distributed system involving several universities and research institu- tions all over the country, and then the Soviet Union fol- lowed suit. A third world data center was established for Europe and Japan. Multiple data sets in different parts of 03_Korsmo_pg023-034_Poles.indd 2903_Korsmo_pg023-034_Poles.indd 29 11/17/08 8:38:19 AM11/17/08 8:38:19 AM 30 SMITHSONIAN AT THE POLES / KORSMO FIGURE 1. The Oceans, IGY Poster. National Academy of Sciences, 1958. 03_Korsmo_pg023-034_Poles.indd 3003_Korsmo_pg023-034_Poles.indd 30 11/17/08 8:38:20 AM11/17/08 8:38:20 AM POLICY PROCESS AND THE INTERNATIONAL GEOPHYSICAL YEAR 31 the world were encouraged to ensure against catastrophic destruction of a single center and to make the data acces- sible to researchers in different parts of the world. While the national IGY committees were responsible for delivering timely and quality data, the world data cen- ters were responsible for the safekeeping, reproduction, cataloging, and accessibility of the data. Anyone engaged in research was to be given access. If you could get your- self into the host country and up to the door of the data center, you could not be turned away (P. Hart, personal communication). We know there were gaps, and the limits of East? West cooperation became immediately apparent in the satellite program. However, the rules of the game were in place, establishing norms of behavior that lasted well beyond the IGY. Finally, the IGY publications included but went much further than peer reviewed scientifi c journals. The Annals of the IGY, 48 volumes published by Pergamon Press be- tween 1959 and 1970 (International Council of Scientifi c Unions, 1959; Fleagle, 1994:170), record not only the re- sults of the research, but also the process of developing the IGY? the international meetings, the world data center guidelines, and the resolutions. This was a self-conscious documentation effort, and we are still benefi ting from that today. We have journalists? accounts such as Walter Sulli- van?s Assault on the Unknown (1961). Upon completion of the IGY research, Sullivan was given virtually unfet- tered access to the IGY fi eld projects and documentation. The establishment of the world data centers and the constant encouragement by the U.S. National Committee for researchers to publish and share their results in many forms demonstrated to many audiences the ability of sci- ence organizations to live up to their commitments. The world data centers, which continue to function today, were instrumental in continuing the legacy of the IGY: coordi- nated, international science to understand the interactions of atmospheric, oceanic, and terrestrial processes. While entrepreneurs initiated schemes and linked solutions to problems, the thousands of people who carried out the work and contributed results gave credibility to the geo- physical sciences, both in the United States and in other countries. Suffi cient trust had been established to begin the demilitarization of geophysics. CONCLUSION The IGY can teach us something about both process and results. The ways in which Berkner, Chapman, Odi- shaw, and many others pursued the activities of agenda- setting, linking solutions to problems, and establishing a pattern of credible commitments, turned a casual conversa- tion among a few experts into a whirlwind series of interna- tional expeditions and experiments. There are many other ways of looking at the IGY. Elsewhere, I have compared my approach to the story of the blind men and the elephant. After they felt different parts of the animal, they each pro- claimed the elephant looked like a tree trunk, a snake, and a fan (Korsmo, 2007a). The policy sciences are useful in deciding what rules of decision-making and allocation of resources work the best for different types of projects in different contexts. The context I chose here was primarily based in the United States, but it would be quite valuable to compare both current and historical science policy evolu- tion in other countries during the time of the IGY. This symposium is an important step in the documen- tation of our present efforts in the International Polar Year 2007? 2008. Due to the precedents set by the IGY and its successors, such as the International Years of the Quiet Sun (1964? 1965) and the Upper Mantle Project (1962? 1970), we have international frameworks for scientifi c coopera- tion as well as a history of data sharing and long-term environmental observations. While the IGY did not include social sciences, it sup- ported a surprising amount of geography and natural his- tory associated with the study of ice sheets and mountain glaciers. The umbrella was large enough to include these projects and contribute to our knowledge of alpine and Arctic ecosystems. The glaciology program of the IGY was a bridge between the physical and biological sciences, paving the way for programs such as UNESCO?s Inter- national Hydrological Decade, 1965? 1974 (Kasser 1967; Muller 1970). In a similar fashion, the social and human studies included in the present International Polar Year 2007? 2008 are a bridge between science and policy. Their inclusion (e.g., Krupnik et al., 2005) provides even more opportunity for self-refl ection and evaluation of what les- sons and legacies we will leave for the future. ACKNOWLEDGMENTS I am grateful to several IGY 1957? 1958 participants, including Pembroke Hart and Phillip Mange, for their insights and recollections. I also wish to thank the ar- chivists and staff of the following institutions for their assistance in locating and accessing the archival and oral history collections used in this research: Sydney Chap- man Papers, University of Alaska Fairbanks, Elmer E. Rasmuson Library, Alaska and Polar Regions Collec- tions; Nathaniel C. Gerson Papers and Alan T. Waterman 03_Korsmo_pg023-034_Poles.indd 3103_Korsmo_pg023-034_Poles.indd 31 11/17/08 8:38:24 AM11/17/08 8:38:24 AM 32 SMITHSONIAN AT THE POLES / KORSMO Papers, U.S. Library of Congress, Manuscripts Division; IGY Collection at the U.S. National Academy of Sciences Archives; Oral History Collection, Niels Bohr Library and Archives, Center for History of Physics, American Institute of Physics; Polar Archival Program, Ohio State University Libraries; and U.S. Research and Development Board Collection, Record Group 330, Entry 341, U.S. National Archives and Records Administration. This ma- terial is based on work supported by the National Science Foundation (NSF) while I was working at the foundation. Any opinions, fi ndings, and conclusions expressed in this article are those of the author and do not necessarily re- fl ect the views of NSF. APPENDIX 1 THE CSAGI BUREAU MEMBERS AND REPORTERS, IPY 1957? 1958 The following list demonstrates the breadth of disci- plines and expertise represented among the international IGY leadership (Nicolet, 1984:314). CSAGI Bureau S. Chapman, President (UK) L. Berkner, Vice President (USA) M. Nicolet, Secretary General (Belgium) J. Coulomb, Member (France) V. Beloussov, Member (USSR) CSAGI Discipline Reporters World Days (when intensive measurements would be taken)? A. H. Shapley (USA) Meteorology? J. Van Mieghem (Belgium) Geomagnetism? V. Laursen (Denmark) Aurora and Airglow? S. Chapman (UK), with F. Roach and C. Elvey (USA) Ionosphere? W. J. G. Beynon (UK) Solar Activity? Y. Ohman (Sweden) Cosmic Rays? J. A. Simpson (USA) Longitudes and Latitudes? J. Danjon (France) Glaciology? J. M. Wordie (UK) Oceanography? G. Laclav?re (France) Rockets and Satellites? L. V. Berkner (USA) Seismology? V. V. Beloussov (USSR) Gravimetry? P. Lejay (France) Nuclear Radiation? M. Nicolet (Belgium) APPENDIX 2 MEMBERS OF THE FIRST U.S. NATIONAL COMMITTEE FOR IPY 1957? 1958 The following list of U.S. IGY leadership shows a mix- ture of disciplinary and professional society representation and the direct involvement of U.S. government agencies (Atwood, 1952). Chair J. Kaplan, representing the U.S. National Committee of the International Union of Geodesy and Geophysics Members L. H. Adams, Geophysical Laboratory, Carnegie Institu- tion of Washington Henry Booker, International Scientifi c Radio Union (URSI) Lyman W. Briggs, National Geographic Society G. M. Clemence, U.S. Naval Observatory C. T. Elvey, Geophysical Institute, University of Alaska John A. Fleming, American Geophysical Union Nathaniel C. Gerson, Cambridge Research Directorate, U.S. Air Force Paul Klopsteg, National Science Foundation F. W. Reichelderfer, U.S. Weather Bureau and World Me- teorological Organization Elliot B. Roberts, U.S. Coast and Geodetic Survey Alan H. Shapley, U.S. Bureau of Standards Paul A. Siple, representing U.S. National Committee of the International Geography Union, Association of American Geographers, and General Staff, U.S. Army Otto Struve, representing U.S. National Committee of the International Astronomy Union Merle Tuve, Department of Terrestrial Magnetism, Carn- egie Institution of Washington Lincoln Washburn, Snow, Ice, and Permafrost Research Establishment, U.S. Corps of Engineers, and the Arctic In- stitute of North America Ex-offi cio Members Wallace W. Atwood Jr., Director, Offi ce of International Relations, National Academy of Sciences-National Re- search Council Lloyd V. Berkner, Member, Special Committee for the In- ternational Geophysical Year J. Wallace Joyce, Deputy Science Advisor, U.S. Depart- ment of State 03_Korsmo_pg023-034_Poles.indd 3203_Korsmo_pg023-034_Poles.indd 32 11/17/08 8:38:25 AM11/17/08 8:38:25 AM POLICY PROCESS AND THE INTERNATIONAL GEOPHYSICAL YEAR 33 NOTE 1. In a similar fashion, the planners of the 2007? 2008 Interna- tional Polar Year came up with a framework that established criteria for participation. This not only provided guidance for researchers who were considering whether to write proposals, but also the existence of criteria conveyed to a broader audience, including policy-makers at the national level, the seriousness and purposefulness of the upcoming science cam- paign (Rapley et al., 2004). LITERATURE CITED Ahlmann, H. W. 1946. Glaciological Methods. Polar Record, 4(31): 315? 319. American Institute of Physics. 2000. Finding Aid to the James A. Van Al- len Papers, 1938? 1990. University of Iowa, Iowa City. http://www. aip.org/ history/ ead/ 19990077.html (accessed 18 September 2007). Atwood, W. W., Jr. 1952. Letter to Chairmen of U.S. National Commit- tees of International Scientifi c Unions, 9 December 1952. In Folder ?Organization USNC 1952.? National Academy of Sciences, IGY Archive, Washington, D.C. Baker, F. W. G. 1982. A Century of International Interdisciplinary Co- operation. Interdisciplinary Science Reviews, 7(4): 270? 282. Beckman, A. O. 1956. ?A Businessman?s View on the ?Failure? of Educa- tion.? U.S. News and World Report, 30 November 1956, 83? 89. Bentley, C. R. 1964. ?The Structure of Antarctica and Its Ice Cover.? In Research in Geophysics, ed. H. Odishaw, vol. 2, pp. 335? 389. Cambridge, Mass.: MIT Press. Beynon, W. J. G. 1975. U.R.S.I. and the Early History of the Ionosphere. Philosophical Transactions of the Royal Society of London. Series A, Mathematical and Physical Sciences, 280: 47? 55. Bullis, H. 1973. The Political Legacy of the International Geophysical Year. Washington, D.C.: U.S. Government Printing Offi ce. Bush, V. 1945. Science? The Endless Frontier. Washington, D.C.: Na- tional Science Foundation. [Reprinted in 1990] Chapman, S. 1955. Letter to Merle Tuve, 8 January. Box 53, Folder 103, ?Data Processing? in ?Sydney Chapman Papers.? Alaska and Po- lar Regions Collections, Elmer E. Rasmuson Library, University of Alaska, Fairbanks. Chapman, S. 1960. From Polar Years to Geophysical Year. Studia geo- physica et geodaetica, 4: 313? 324. Cusack, T. R., T. Iversen, and D. Soskice. 2007. Economic Interests and the Origins of Electoral Systems. American Political Science Re- view, 101(3): 373? 391. Day, D. A. 2000. ?Cover Stories and Hidden Agendas: Early Ameri- can Space and National Security Policy.? In Reconsidering Sput- nik: Forty Years Since the Soviet Satellite, ed. R. D. Launius, J. M. Logsdon, and R. W. Smith, pp. 161? 195. Amsterdam: Harwood Academic Publishers. Fleagle, R. G. 1994. Global Environmental Change: Interactions of Science, Policy, and Politics in the United States. Westport, Conn.: Praeger. Gerson, N. C. 1953. ?1st USNC Meeting, 27 March 1953.? Box 8, Folder 1, ?N. C. Gerson Papers.? Manuscripts Division, Library of Congress, Washington, D.C. Good, G. A. 2000. ?Biographical Note.? In Guide to the Sydney Chap- man Papers. Alaska and Polar Regions Collections, Elmer E. Ras- muson Library, University of Alaska Fairbanks. http://nwda-db. wsulibs.wsu.edu/ fi ndaid/ ark:/ 80444/ xv48580#bioghistID (accessed 18 September 2007). Hall, R. C. 1997. The Truth about Overfl ights. Quarterly Journal of Military History, 9(3): 24? 39. International Council of Scientifi c Unions. 1959. Annals of the Interna- tional Geophysical Year, Vol. 1. New York: Pergamon Press. Kasser, P. 1967. Fluctuations of Glaciers, 1959? 1965. Technical Papers in Hydrology, 1. Paris: UNESCO (ICSI). Kevles, D. J. 1990. Cold War and Hot Physics: Science, Security, and the American State, 1945? 56. Historical Studies of the Physical Sci- ences, 20(2): 238? 264. Kingdon, J. W. 1995. Agendas, Alternatives, and Public Policies. 2nd ed. New York: Addison-Wesley Educational Publishers. Kirwan, L. P., C. M. Mannerfelt, C. G. Rossby, and V. Schytt. 1949. Glaciers and Climatology: Hans W:son Ahlmann?s Contribution. Geografi ska Annaler, 31: 11? 20. Korsmo, F. L. 2004. Shaping Up Planet Earth: The International Geo- physical Year (1957? 1958) and Communicating Science through Print and Film Media. Science Communication, 26(2):162? 187. ???. 2007a. The International Geophysical Year of 1957 to 1958. Sci- ence, People and Politics 2 (January 2007). http://www.gavaghan- communications.com/korsmoigy.html (accessed 14 September 2007). ???. 2007b. The Genesis of the International Geophysical Year. Phys- ics Today, 60(7): 38? 43. Korsmo, F. L., and Sfraga, M. P. 2003. From Interwar to Cold War: Sell- ing Field Science in the United States, 1920s? 1950s. Earth Sciences History, 22(1): 55? 78. Krupnik, I., M. Bravo, Y. Csonka, G. Hovelsrud-Broda, L. M?ller-Wille, B. Poppel, P. Schweitzer, and S. S?rlin. 2005. Social Sciences and Humanities in the International Polar Year 2007? 2008: An Inte- grating Mission. Arctic, 58(1): 91? 97. M?ller, F., ed. 1970. Perennial Ice and Snow Masses: A Guide for Com- pilation and Assemblage of Data for a World Inventory; Variations of Existing Glaciers: A Guide to International Practices for Their Measurement. Technical Papers in Hydrology, 5. Paris: UNESCO (ICSI). National Academy of Sciences. 1958. Planet Earth: The Mystery with 100,000 Clues. Washington, D.C.: National Academy of Sciences. Nicolet, M. 1984. The International Geophysical Year (1957? 1958): Great Achievements and Minor Obstacles. GeoJournal, 8(4): 303? 320. Needell, A. 2000. Science, Cold War, and the American State: Lloyd V. Berkner and the Balance of Professional Ideals. Washington, D.C.: Smithsonian Institution and Harwood Academic. North, D. C., and B. R. Weingast. 1989. Constitutions and Commit- ment: The Evolution of Institutions Governing Public Choice in Seventeenth-Century England. The Journal of Economic History, 49(4): 803? 832. Rapley, C., R. Bell, I. Allison, R. Bindschadler, G. Casassa, S. Chown, G. Duhaime, V. Kotlyakov, M. Kuhn, O. Orheim, P. C. Pandey, H. K. Petersen, H. Schalke, W. Janoschek, E. Sarukhanian, Z. Zhang. 2004. A Framework for the International Polar Year. ICSU IPY 2007? 2008 Planning Group. http://216.70.123.96/ images/ uploads/ framework.pdf (accessed 19 September 2007). Riker, W. 1962. The Theory of Political Coalitions. New Haven, Conn.: Yale University Press. Rossby, C. G. 1947. Letter to Executive Secretary, Joint Research and Development Board, 21 April 1947. Research and Development Board, Record Group 330, Entry 341, Box 452, Folder 2. National Archives and Record Administration, College Park, Md. Sabatier, P. A., and H. C. Jenkins-Smith. 1999. ?The Advocacy Coalition Framework: An Assessment.? In Theories of the Policy Process, ed. P. A. Sabatier, pp. 117? 166. Boulder, Colo.: Westview Press. Siple, P. 1948. ?Memorandum to Robert B. Simpson, 14 April 1948,? p. 4. Record Group 330, Entry 341, Box 452, Folder 1. U.S. Na- tional Archives and Records Administration, College Park, Md. Sullivan, W. 1961. Assault on the Unknown: The International Geophys- ical Year. New York: McGraw-Hill. 03_Korsmo_pg023-034_Poles.indd 3303_Korsmo_pg023-034_Poles.indd 33 11/17/08 8:38:25 AM11/17/08 8:38:25 AM 34 SMITHSONIAN AT THE POLES / KORSMO U.S. National Archives and Records Administration, n.d. ?Finding Aid? to Record Group 330, Entry 341, ?Offi ce of the Secretary of De- fense, Research and Development Board.? U.S. National Archives and Records Administration, College Park, Md. U.S. Research and Development Board. 1953. Memorandum from the Board, 23 May, in General Correspondence, Box 33. ?Alan T. Waterman Collection.? Library of Congress, Washington, D.C. U.S. National Committee for the IGY. 1960. Press release, 17 Novem- ber 1960. National Academy of Sciences IGY Archive, Offi ce of Information, Series 12, Planet Earth Films Chron. File. National Academy of Sciences, Washington, D.C. U.S. National Security Council. 1957. Statement of Policy by the National Security Council on Antarctica. NSC 5715/1. 29 June. Contained in NSC Series, Policy Papers Subseries, Eisenhower Library. Abilene, Kans. Van Allen, J. A. 1997. Interview with B. Shoemaker, November 18, 1997. Transcript. Oral History Collection, Ohio State University, Columbus. ???. 1998. Genesis of the International Geophysical Year. The Polar Times, 2(11):5. 03_Korsmo_pg023-034_Poles.indd 3403_Korsmo_pg023-034_Poles.indd 34 11/17/08 8:38:26 AM11/17/08 8:38:26 AM ABSTRACT. In July 1966, the 89th Congress (H.R. 6125) laid out the charge defi ning the new Smithsonian National Air and Space Museum: to ? memorialize the national development of aviation and space fl ight; collect, preserve, and display aeronautical and space fl ight equipment of historical interest and signifi cance; serve as a repository for scientifi c equipment and data pertaining to the development of aviation and space fl ight; and provide educational material for the historical study of aviation and space fl ight.? Under this umbrella statement, the Museum has been actively collecting artifacts and documentary evidence in the area of the earth and space sciences, as well as in astronomy, that helps to preserve the social, cultural, intellectual, and material legacy of the enter- prise. The paper examines the holdings pertaining to the IGY era (1957? 1960) presently in the NASM collection. It discusses how some of these items were identifi ed, selected, and collected, as a means of offering a preliminary appraisal of the historical value of the collection. It highlights a suite of objects built by James Van Allen?s Iowa group and discusses their historical signifi cance. THE IGY The International Geophysical Year (IGY) of 1957? 1958 was conceptual- ized at a small dinner party in April 1950, held at the home of James A. Van Allen in Silver Spring, Maryland. As Walter Sullivan recorded at the time, and as Fae Korsmo and many others have reminded us more recently (Sullivan, 1961; Korsmo, 2007; this volume), out of this meeting grew a plan to coordinate ob- servations relevant to the geosciences over all parts of the globe, and, for the fi rst time, conduct signifi cant soundings of the upper reaches of the earth?s at- mosphere and ionosphere. Considering that three members of the party, notably Van Allen, Lloyd Berkner, and Sydney Chapman, later to become key players in IGY, were primarily concerned with studying the ionosphere, it is not surpris- ing that they organized the means to pursue its global characteristics using all available technologies (Needell, 2000). Their plan was aided and abetted by Cold War priorities for developing the capabilities of space fl ight to aid global reconnaissance, and in fact became driven by those priorities, modifi ed in com- plex ways by the foreign policy and national security strategies of the major participating nations (McDougall, 1985; Bulkeley, 1991). David DeVorkin, National Air and Space Mu- seum, Smithsonian Institution, P.O. Box 37012, MRC 311, Washington, DC 20013-7012, USA (devorkind@si.edu). Accepted 29 May 2008. Preserving the Origins of the Space Age: The Material Legacy of the International Geophysical Year (1957? 1958) at the National Air and Space Museum David H. DeVorkin 04_DeVorkin_pg035-048_Poles.indd35 3504_DeVorkin_pg035-048_Poles.indd35 35 11/17/08 8:35:28 AM11/17/08 8:35:28 AM 36 SMITHSONIAN AT THE POLES / De VORKIN Out of this complex mixture of scientifi c and national security priorities, both the Soviet Union and the United States announced plans to orbit artifi cial satellites during the IGY, and both made good on their promise, though in a manner, and especially an order, that surprised and deeply disturbed a large portion of world?s media and propelled a space race between the two superpowers that fuelled the fi rst decade of what has been called the Space Age. A substantial historical literature exists recounting the IGY and the origins of the space age (Pisano and Lewis, 1988; Marson and Terner, 1963; McDougall, 1985) Our purpose here is not to recount this history nor to delineate the Smithsonian Institution?s participation in space research, but rather to describe holdings at the National Air and Space Museum (NASM) pertaining to space fl ight activities dur- ing the International Geophysical Year (1957? 1958). This paper discusses specifi cally how the Smithsonian National Air and Space Museum (NASM) has participated in pre- serving the material heritage of this legacy? a legacy that might be seen someday as a major factor leading to the exis- tence of the Museum itself. We begin by situating the act of cultural preservation within the mission of the Smithsonian, and then conclude with an assessment of efforts to preserve the material legacy of the IGY. ON MUSEUM PRESERVATION Institutions like the NASM collect for a variety of purposes, both immediate and long term. One can only speculate about why, precisely, a material legacy will be important for our descendents, say 400 years from now; whether they are specialists in science and its history, or whether they are educated and inquisitive nonspecialists. Will they think well of us for making the effort to pre- serve a material legacy, one that can be ?read? without the intervention of media-specifi c technologies? Or will they possess technologies undreamt of today for seeking out the answers to historical questions that transcend the ma- terial legacy, and regard our efforts as ultimately futile? The mission, defi ned by legislation that brought the NASM into existence, claims that the Institution has a re- sponsibility to ?memorialize the national development of aviation and space fl ight? (U.S. House of Representatives, 1946). That is what we indeed do, and as I have argued elsewhere, the survival of a physical artifact will, in and of itself, help to stimulate questions about our times some- day, and may even, conceivably, help to answer questions about our lives and times (DeVorkin, 2006a; 2006b). As more than one observer has noted recently, commenting about the signifi cance of objects displayed in museums and the motives for the curators to put them on display; ?Their presence there is the message. . . . It is still all about visibility.? (Kennicott, 2007:C1) From the beginning, NASM?s charge has been to ?serve as the repository for, preserve, and display aero- nautical and space fl ight equipment and data of historical interest and signifi cance to the progress of aviation and space fl ight, and provide educational material for the his- torical study of aviation and space fl ight and their technol- ogies? (U.S. House of Representatives, 1946) Since this is a formal process with oversight, NASM curators periodi- cally create and review collections plans that rationalize the effort. But curators are also keenly aware of the fact that they are, in some way, making choices and hence are fi ltering history. After all, taking the existentialist?s point of view, as Oxford?s Jim Bennett and others have observed from time to time ?museum collections . . . show you not what there was but what was collected.? (Bennett, 2004), This statement is a simple fact of life, of means, motives, and of circumstance. Unlike the natural history disciplines, whose collections stand at the very core of their research interests, and in fact defi ne them, forming the data banks from which they ask questions and draw conclusions, col- lections of space history refl ect something rather different. They refl ect cultural and institutional needs to preserve the material heritage of ourselves, a very recent past still very much alive in its human participants and in its institutions, and of which we are part, our contemporary national heri- tage. Deciding what to collect and preserve, then, involves as much issues of a symbolic nature, the need to memo- rialize, as it does intellectual issues relating to any disci- plinary goals, past, present, or future, and so the questions historians ask about culture transcend the specimens they preserve. Collections are not comprehensive of their culture and are the result of choices, personal preferences, biases, and both political and fi nancial limitations. So what is it that institutions like museums do when they collect and preserve? The economic anthropologist tells us that the act of collection by institutions is a for- mal method of removing objects from the commodity sphere? the sphere of use, speculation, and trade? and placing them into a singularized and sacralized sphere (Kopytoff, 1986). This is defi nitely what we try to do at the Smithsonian and it is refl ected in our Collections Ra- tionales, our arguments over what to collect. Indeed, over the past years, some curators have repeatedly worried that if we do not maintain control of objects deemed to be of historical value coming out of the Nation?s space program, relics bought and paid for by taxpayers, they will be sold 04_DeVorkin_pg035-048_Poles.indd36 3604_DeVorkin_pg035-048_Poles.indd36 36 11/17/08 8:35:28 AM11/17/08 8:35:28 AM PRESERVING THE ORIGINS OF THE SPACE AGE 37 as excess property and become commodities for specula- tion by collectors and agents. In more recent times, as our ability to collect has met serious fi nancial, personnel, and storage limitations, this concern has diminished somewhat and there is now healthy consideration of establishing a means to distribute the responsibility of preservation. This includes establishing a ?national strategy? of shar- ing the responsibility for preserving the heritage among many institutions. But ultimately, we generally accept the view that what it is we are doing by collecting is making this material heritage accessible to future generations in a manner that will stimulate interest and remembrance of an historical era or event. A clear symptom of this rationale for sacralization and protection from the commodity sphere comes from a unique agreement the Smithsonian Institution maintains with NASA, the ?NASA/NASM Transfer Agreement? (Agreement, 1967). This document asserts that any object on NASA?s inventories that is deemed by a select committee of NASA program managers and specialists to be historic and excess to present agency needs must be offered fi rst to the National Air and Space Museum for its collection. The NASM will then deliberate and decide upon collection. If it agrees, the object is transferred to the Smithsonian in- ventory. If not, it goes on the normal ?excess property? listings and can be transferred to other agencies, or sold to the public. It is the existence of this agreement that gives the Space History collection at NASM its special respon- sibility. Even though the IGY-era collections predate the agreement, many came to the Museum as a result of its existence, well after the close of the era. There is yet another aspect of collecting in Space His- tory that warrants attention here that adds to its unique character and will help us evaluate our IGY-era collections. Many of the most important objects, those responsible for the actual science performed, are not available for collec- tion and never will be. They were launched, and were either consumed through use or by re-entry, or are now in orbits that make them inaccessible. What we can collect, there- fore, are surrogates for the ?real thing.? They may be very close in form and function, like fl ight backups, but they are not the actual objects that made the historic observations or performed the historic feats, such as the fi rst soft land- ing on the moon, Mars, Titan, or an asteroid. There are exceptions, of course, such as the panoramic camera from Surveyor III that was returned to Earth by Apollo 12 astro- nauts (NASM Catalogue number I19900169001; hereafter, just the alphanumerical code will be used: ?I? for incoming loan, ?A? for accession). There are objects returned from Shuttle missions, as well as objects that returned to earth by design, like the particle collectors aboard Stardust, and interplanetary probes that were launched and might some- day be captured and returned, like the third International Sun-Earth Explorer (ISEE-3). In addition to its special relationship with NASA, and its unique responsibilities, NASM must still justify what it collects both internally and externally. Individu- als rarely rationalize why they collect what they collect but institutions, especially public ones, must provide clear and cogent rationalizations in order to gain the support to identify, collect, and preserve. One need only consider the large costs involved in collection and preservation, and the long-term commitment an institution or a culture is willing to make in supporting such efforts. Thus, given this mandate, and limitations, how representative is ?what was collected? to ?what there was?? THE TIME BOUNDARIES OF THE IGY LEGACY The legacy of the International Geophysical Year (IGY) of 1957? 1958 predates NASA, of course, though many of the objects that one can describe as belonging to the IGY era came from NASA, which inherited the legacy upon its formation in August 1958 and retained much of it for years at its visitor centers and in storage. We can roughly limit the IGY-era legacy in space by establishing its begin- ning as the material legacy growing out of planning for the IGY since the early 1950s (see Korsmo, this volume), to the launch of Explorer VII in October 1959, Vanguard III in September 1959, and two Discoverer launches (VII and VIII) through 20 November 1959 (Green and Lomask, 1970). Explorer VII was the last Army Ballistic Missile Agency (ABMA) satellite and was transferred to NASA. It represents the end of the legacy started by Sputnik 1 and Explorer 1 and the IGY context, even though transfer of all programs to NASA took place with NASA?s creation in the fall of 1958 via the National Aeronautics and Space Act. The major American programs linked to the IGY era include Vanguard (I? III), Explorer (I? VII), Pioneer (I? IV), and the military programs known as Project SCORE and Discoverer (I? VIII). Russian programs included Sputnik (I? III) and Luna (I? III). THE NASM COLLECTION More than 200 objects in the national collection can be associated with IGY-era space-related activities. These reside in several NASM sub-collections, including rocketry 04_DeVorkin_pg035-048_Poles.indd37 3704_DeVorkin_pg035-048_Poles.indd37 37 11/17/08 8:35:28 AM11/17/08 8:35:28 AM 38 SMITHSONIAN AT THE POLES / De VORKIN and propulsion, the space sciences, memorabilia, interna- tional, and social and cultural collections. There are partial- scale and full-scale models, replicas, engineering models, components, medals, badges, and ephemera (collectibles). In this review, we consider only the artifact collections, not fl at materials in our library and archives, including mono- graphs, serials, technical publications, print and fi lm re- sources, as well as extensive manuscript collections or oral histories (see http:// www.siris.si.edu/ and http://www.nasm .si.edu/ research/ arch/ collections.cfm). IGY-RELATED COLLECTIONS: PREPARING FOR THE IGY In addition to numerous examples of early sound- ing rocket payloads for both atmospheric and space re- search? such as ultraviolet spectrographs, X-ray detectors, magnetometers, varieties of halogen quenched particle fl ux counters, mass spectrometers, temperature and pressure sensors, and cameras built for V-2, Viking, Aerobee and ARCAS fl ights from the late 1940s through the 1950s? the NASM collection preserves objects intended specifi cally for use during the IGY, such as a visual Project Moonwatch telescope (A19860036000) (Figure 1) and the fi rst Baker- Nunn satellite tracking camera (A19840406000) mounted in Arizona. The most curious object, symbolic of the aspi- rations of S. Fred Singer, one of the original members of the group that conceived the IGY, is MOUSE (Minimal Orbital Unmanned Satellite, Earth), a full-scale design concept model for an artifi cial satellite (A19731670000). It car- ries two Geiger counters for cosmic-ray studies, photocells, telemetry electronics, and a rudimentary magnetic data storage element (Figure 2). The rocketry collection has ex- amples of small sounding rockets derived from barrage ord- nance technology, such as the Loki-Dart (A19750183000; 0184000), and various combinations that were used in the 1950s for atmospheric measurements and as payloads un- der Skyhook balloons for extreme high-altitude soundings by the University of Iowa and by the Navy. IGY-RELATED COLLECTIONS: SPUTNIK In addition to one of the fi rst full-scale models of Sput- nik 1 on loan to the Smithsonian from the Soviet Union/ Russia (I19900388002), as well as the original electrical arming pin removed from the fl ight unit just before launch (I19971143001), the Museum holds six objects relating to the fl ight, all in the collectible category and ranging from a cigarette lighter and commemorative pin to med- als and a music box. There are no holdings relating to, or informing, the technical characteristics of any of the early FIGURE 1. Project Moonwatch telescope (A19860036000), pre- served and on display at the NASM?s Hazy Center. Amateurs and commercial organizations alike built thousands of instruments of this type; they exist in many forms. The Smithsonian Astrophysical Observatory created and coordinated the Moonwatch program to produce preliminary orbital elements of the fi rst satellites. (NASM photograph) FIGURE 2. S. Fred Singer?s full-scale concept model for an artifi cial satellite (A19731670000). (Eric Long photograph, NASM) 04_DeVorkin_pg035-048_Poles.indd38 3804_DeVorkin_pg035-048_Poles.indd38 38 11/17/08 8:35:29 AM11/17/08 8:35:29 AM PRESERVING THE ORIGINS OF THE SPACE AGE 39 Sputniks. There is nothing in the collection pertaining to Project Luna. IGY-RELATED COLLECTIONS: PROJECT VANGUARD The NASM collection has seven objects identifi ed as Vanguard 1 backup models, test models, replicas, and dis- play models (A19580115000 through A19830244000). Among these is the original test vehicle TV-3 (Figure 3), which was recovered after the launch vehicle crashed onto the launchpad on 6 December 1957 (A19761857000). The object was acquired from John P. Hagen (1908? 1990), the former Project Vanguard manager, in the spring of 1971 and placed on exhibit in the Smithsonian?s Arts & Indus- tries Building. After NASM opened in July 1976, visitors encountered it in the outstretched hand of an unhappy and concerned 12-foot tall ?Uncle Sam.? It now resides in a case near the Museum?s Vanguard rocket, a TV-2BU (Figure 4, center) that had been prepared for launch by the Martin Company on 3 September 1957 but was delayed and then cancelled. Vanguard 1 was launched on a near duplicate rocket; the markings on the NASM version were changed to be identical to those of the fl ight vehicle by the Naval Research Laboratory, which then donated the rocket to the Smithsonian Institution in 1958 (A19580114000). There are also three elements of various stages of the FIGURE 3. Examination of the original TV-3 satellite in March 2008, by members of the Naval Research Lab team who designed and built it, on the eve of the fi ftieth anniversary of the fi rst successful Vanguard fl ight. The object was opened to allow inspection for identifi cation of components, search for undocumented experiments, and to assess its state of preservation. Martin Votaw, left, Roger Easton, right. (Photograph courtesy Judith Pargamin) 04_DeVorkin_pg035-048_Poles.indd39 3904_DeVorkin_pg035-048_Poles.indd39 39 11/17/08 8:35:31 AM11/17/08 8:35:31 AM 40 SMITHSONIAN AT THE POLES / De VORKIN Vanguard propulsion system in the collection, as well as a set of 18 electrical and electronic radio instruments used in a Vanguard Minitrack station (A19761036000). There is one instrumented replica of the Vanguard Lyman Alpha satellite, also called SLV-1, that failed to orbit in May 1958, and two versions of Vanguard III, including Vanguard 3, also called Magne-Ray Satellite, and fi nally the Vanguard Magnetometer satellite that did fl y, desig- nated SLV-5 or Vanguard 3a (A19751413000; 1407000; 1412000). It was placed in orbit in December 1959 and was equipped with two X-ray detectors and micrometeor- oid detectors. The instrumented Vanguards do preserve many of the technical parameters of the early fl ight objects. They were built generally by the same people, using the same jigs and materials, which produced the fl ight objects at the Naval Research Laboratory. Some of the craft have Lyman alpha and X-ray detectors identical to, or very similar to those used by NRL scientists in sounding rocket fl ights through- out the 1950s (DeVorkin, 1996). IGY-RELATED COLLECTIONS: EXPLORER There are more than two dozen objects in the collec- tion relating to some form of Explorer satellite between Explorers I through VII, and dozens more UV and X-Ray detectors identical to those fl own by groups at NRL, Iowa and elsewhere. This collection contains objects with a high degree of historical accuracy and signifi cance, and pre- serves some of the most detailed technical characteristics of the fi rst fl ight objects. Of great historical signifi cance is a suite of objects recently acquired from George H. Lud- wig, one of James Van Allen?s graduate students at the time the Iowa group became engaged in preparing for the fi rst Explorers. The primary object Dr. Ludwig donated was built as an engineering model for the payload for the fi rst fully instrumented Vanguard fl ight (Figure 5), and then became the template for the redesign effort at the Jet Propulsion Laboratory after Army Ordnance was given the green light to proceed with a launch after the failure of the Vanguard TV-3 launch (A20060086000). As both Van Allen and Ludwig have noted in various recollections (Van Allen, 1983:55? 57), the package was designed for a 20-inch Vanguard sphere but was within parameters easily adaptable to a 6-inch diameter cylindrical chamber com- patible with the dimensions of the scaled-down Sergeant solid rocket of the sort that ABMA had been placing on Jupiter test fl ights. So when a fl ight on a Jupiter-C became possible in the wake of the Vanguard failure, and after Von Braun promised Eisenhower that the Army could or- FIGURE 4. Vanguard launch vehicle TV-2BU on display at NASM in the ?Missile Pit.? (NASM photograph by Eric Long) 04_DeVorkin_pg035-048_Poles.indd40 4004_DeVorkin_pg035-048_Poles.indd40 40 11/17/08 8:35:35 AM11/17/08 8:35:35 AM PRESERVING THE ORIGINS OF THE SPACE AGE 41 bit something useful within 90 days, Ludwig packed his bags and his family and brought their Vanguard payload to JPL for modifi cation into what would be called at fi rst ?Deal 1.? The object in the collection includes electronic and mechanical elements of the initial ?Deal? payloads designed and built by the University of Iowa. In addition, Ludwig donated versions of the separate electronic com- ponents before they would have been ?potted? or elec- tronically sealed for fl ight. All of these components had been in Ludwig?s possession since they were built in the late 1950s, with the exception of short intervals when the components were at JPL under his care. Their provenance, in other words, is unquestioned, and he has provided ex- tensive documentation attesting to their historical role in the early Explorer series. Possibly the most signifi cant corrective to our docu- mentation of IGY-era artifacts in the NASM collection came as a result of Ludwig?s assistance. In 1961, the Jet Propulsion Laboratory transferred what it claimed was a fully instrumented fl ight spare of Explorer 1 to the Smith- sonian Institution (A19620034000). Attached to an empty fourth-stage Sergeant rocket, it was initially displayed in the Smithsonian?s Arts & Industries building to symbolize the United States? fi rst successful artifi cial satellite. It became a centerpiece of the NASM Milestones gallery upon open- ing in 1976, having toured briefl y just before the opening (Figure 6). In 2005, acting upon an inquiry from Ludwig (2005a), who was then searching out all surviving cosmic- ray Geiger counter detectors inventoried in Van Allen?s Iowa laboratory that supplied the fi rst Explorers, the Collections Management staff of NASM, led by Karl Heinzel, removed the object from display for dismantling (Figure 7), conser- vation evaluation, and inspection. It was empty (Figure 8). Naturally, we reported this fact back to Ludwig, af- ter taking detailed photographs of the interior instrument frame, wiring and markings. The micrometeoroid detector was in place, wrapped around the external shell, but not any of the associated electronics. Nothing from the cosmic ray package survived. But there were clear markings that at one time, the instrument frame had held ?Payload II? as those words appeared in red (Figure 8). Ludwig?s highly detailed documentary record showed, immediately, that this was indeed the fl ight backup for ?Deal 1? that was sent back to Van Allen?s Iowa laboratory for inspection and testing, and then was returned to JPL later in 1958. The FIGURE 5. George Ludwig examining his engineering model for the payload for the fi rst fully instrumented Vanguard fl ight and became the template for the Explorer 1 payload. (NASM photograph by Dave DeVorkin) FIGURE 6. Explorer 1 (A19620034000), initially displayed in the Smithsonian?s Arts & Industries building, became a centerpiece in NASM?s Milestones gallery upon opening in 1976. Opening it for inspection revealed it was a true backup but was devoid of instru- mentation. (NASM photograph) 04_DeVorkin_pg035-048_Poles.indd41 4104_DeVorkin_pg035-048_Poles.indd41 41 11/17/08 8:35:38 AM11/17/08 8:35:38 AM 42 SMITHSONIAN AT THE POLES / De VORKIN cosmic-ray package donated by Ludwig in 2006 was indeed the Vanguard payload that served as template for the fl ight version of Deal 1. Ludwig is not absolutely positive that his donated package was originally inside the spacecraft we display today in Milestones, but it is identical in nature (Ludwig, 1959, 1960, 2005). IGY-RELATED COLLECTIONS: PIONEER Van Allen?s group was also engaged to instrument a se- ries of Pioneer fl ights under Air Force auspices that were aimed at the moon. Although not successful in this goal, three of them managed to detect the complete inner and outer structures of the Earth?s radiation belts, as well as confi rm the profound infl uence that solar activity has on the Earth?s radiation environment. The fi rst Pioneer reached 70,000 miles altitude, less than a third the distance to the Moon, but failed to achieve either orbital or escape velocity and so re-entered the earth?s atmosphere and was destroyed. The Pioneer 1 replica in the collection was reconstructed out of original parts that failed to meet fl ight specifi cations (A19640665000). Two examples of the smaller Pioneer IV are also in the collection. One is a cutaway model showing the instrumentation and housekeeping elements, and the other is a fully instrumented fl ight spare (A19751426000; A19620018000). FIGURE 7. Inspection and conservation evaluation of Explorer 1 (A19620034000) by Matthew Nazarro, specialist at the NASM?s Paul E. Garber Restoration Facility. (NASM photograph) 04_DeVorkin_pg035-048_Poles.indd42 4204_DeVorkin_pg035-048_Poles.indd42 42 11/17/08 8:35:40 AM11/17/08 8:35:40 AM PRESERVING THE ORIGINS OF THE SPACE AGE 43 IGY-RELATED COLLECTIONS: LAUNCH VEHICLES AND SUPPORT EQUIPMENT The NASM Collection boasts probably the strongest collection of IGY-era launch vehicles in the world. As noted above, a full and virtually complete multi-stage Vanguard TV-2BU is preserved, as well as examples of turbo-pumps and engines. In addition, the precursors to its various stages are in the collection, including a full-scale Viking rocket that was prepared for display in the early 1950s using real components from the Glenn L. Martin Company and Reac- tion Motors inventories. A Jupiter-C missile, donated by the Army in 1959, dominates NASM?s ?Missile Pit? (Figure 4, left) and is capped by a complete array of scaled-down Ser- geant rockets in a confi guration identical to that used for the launch of Explorer 1 (A19590068000). Numerous Aerobee and Aerobee-Hi components, along with a complete unit, document the most prolifi c sounding rocket in history. The collection also contains many smaller solid rockets, most with tactical air-to- ground and ground-to-air origins, and even one example of the multistage Farside, a gigantic balloon-launched rocket system created for the Air Force Offi ce of Scientifi c Research for extreme high-altitude non-orbital fl ights in the fall of 1957 (A19680013000). Although there are several Loki-based multistage sys- tems in the collection, we lack a fully articulated ?Rock- oon? system consisting of a Skyhook-balloon, connecting hardware, radio telemetry control, and a small solid- fuelled rocket, such as a Loki or Loki-Dart. These systems, in the hands of James Van Allen?s Iowa team, as well as various NRL groups, carried cosmic-ray detector payloads FIGURE 8. Inspection revealed that Explorer 1 (A19620034000) was empty but that at one time, the instrument frame had held ?Payload II.? (NASM photograph.) 04_DeVorkin_pg035-048_Poles.indd43 4304_DeVorkin_pg035-048_Poles.indd43 43 11/17/08 8:35:43 AM11/17/08 8:35:43 AM 44 SMITHSONIAN AT THE POLES / De VORKIN to high altitudes at a wide range of geographic latitudes during the IGY. IGY-RELATED COLLECTIONS: INDIVIDUALS There is little question today that the one name that will survive from the IGY in historical accounts written in future centuries will be James Van Allen. From a cur- sory analysis of newspaper coverage of the IGY era based upon a Proquest survey conducted by Sam Zeitlin, 2007 NASM Summer intern, Van Allen?s name stands out above all others as most frequently cited or referred to. A sig- nifi cant region of space surrounding the earth has been named for him, the ?Van Allen belts,? a term which at this writing garners more than 78,000 ?hits? in a simple Google search. So it is reasonable to ask: What have we done to preserve the material legacy of James Van Allen at NASM? Van Allen was a Regents? Fellow at the Smithsonian in 1981, spending much of the academic year preparing a personal scientifi c memoir and submitting himself to some 18H11001 hours of oral history interviews by NASM curators and historians (SAOHP, NASM Archives). He also partici- pated in several symposia and seminars (Hanle and Von del Chamberlain, 1981; Mack and DeVorkin, 1982; Van Allen, 1983). He was then planning for the organization of his papers at Iowa, where they would be housed, and engaged NASM staff in an advisory capacity to appraise the collection. Out of this intimate contact, Van Allen eventually donated a small selection of objects that both informs and symbolizes his career. In the early 1990s, he donated a casing from a World War II? era Mark 58 radio proximity fuze for antiaircraft artillery fi re control (Figure 9). The fuze had been partly cut open to display the mi- croelectronic components (A19940233000). He was part of the wartime effort to design, test, and build these fuzes, FIGURE 9. World War II? era Mark 58 radio proximity fuze for anti-aircraft artillery fi re control designed by the Applied Physics Laboratory group that included Van Allen. (NASM photograph) 04_DeVorkin_pg035-048_Poles.indd44 4404_DeVorkin_pg035-048_Poles.indd44 44 11/17/08 8:35:46 AM11/17/08 8:35:46 AM PRESERVING THE ORIGINS OF THE SPACE AGE 45 and Van Allen played a signifi cant role in bringing them into operation through tours with the fl eet in the Pacifi c. The experience and expertise he gained managing his por- tion of this program served him well after the war when he devoted much of his energies to building delicate and com- plex arrays of Geiger counters for rocket fl ights aboard captured German V-2 missiles, then Aerobees, and espe- cially the innovative balloon-launched Loki-Dart systems (Figure 10) he developed at Iowa that subjected payloads to huge accelerations and confi ned quarters reminiscent of the fuze-equipped 5-inch shells (Figure 9). He also do- nated a complete fl ight backup payload for Explorer IV; the fi rst payload designed with knowledge of the existence of the trapped radiation fi eld, and thereby employed new Anton detectors that had smaller cross sections, as well as two small scintillation detectors. Van Allen also asked his Iowa staff to refurbish a plaque bearing a gold-plated fl ight-spare tape recorder that commemorated the fl ight of Explorer III, the fi rst to be able to record and ultimately transmit a continuous record of the radiation fi elds it was encountering, and the fi rst to show unambiguously the presence of the inner regions of trapped radiation. Some 17 objects in the collection preserve the char- acter of the Explorer 1? 4 series and Van Allen?s contribu- tion. Among them is a fl ight spare radio transmitter for Explorer 1, donated by Henry L. Richter, former head of the JPL group that built these units. Richter found that JPL had discarded this unit some years later, and saved it from oblivion. But the surviving mercury-cell batteries in the unit had seriously corroded the overall structure, so Richter removed the corrosive elements, fully docu- mented the process he took to neutralize and restore the structure, and donated the object to the museum in 1997 (A19980115000). Van Allen is not the only pioneer space scientist to be represented in the collection. Objects relating to the efforts of two groups at the Naval Research Laboratory headed by Richard Tousey and Herbert Friedman, over the period starting with V-2 fl ights and lasting through the 1960s and 1970s, have also been preserved and are on display (De- Vorkin, 1992; 1996). GENERAL OBSERVATIONS It should be evident from this brief reconnaissance of IGY-era objects in the NASM collection that, even though our holdings may be impressive, they are not the result of a single rational process or any consistent, premeditated program to preserve IGY history. Some of the objects exist because they were part of the developmental process lead- ing to the fl ight instrument. Others are replicas or facsimi- les created to symbolize the historic event (Figure 11). Most were collected with little or no apparent priority given to engineering, scientifi c or symbolic value, though they had to possess at least one of those qualities. Some objects came to us serendipitously, some due to our intrinsic visibility and centrality to national preservation. Some were collected as the result of judicious inquiry, but not to the extent of spe- cifi cally collecting the IGY era. There were, however, some consistently applied schemes. For instance, we planned out a collection documenting 30 years of electronic ultraviolet and X-ray detector develop- ment by Herbert Friedman?s group at the Naval Research Laboratory (1949? 1980s); there was a similar program trac- ing the evolution of ultraviolet detectors for solar research FIGURE 10. James Van Allen holding a balloon-launched Loki-Dart payload developed at Iowa. ( James A. Van Allen Papers, The Uni- versity of Iowa Libraries, Iowa City, Iowa; c. 1950s) 04_DeVorkin_pg035-048_Poles.indd45 4504_DeVorkin_pg035-048_Poles.indd45 45 11/21/08 3:45:26 PM11/21/08 3:45:26 PM 46 SMITHSONIAN AT THE POLES / De VORKIN with rockets over the same period by members of Richard Tousey?s NRL group, and the efforts of similar teams were devoted to aeronomy and ionospheric physics in the 1950s at NRL, the Applied Physics Laboratory, at the University of Colorado, at the Air Force Cambridge Research labora- tory, and elsewhere (DeVorkin, 1992; 1996; Hirsh, 1983; Schorzman, 1993). Collections arising from these efforts produced a large set of oral and video-histories, as well as a considerable cache of non-record archival material, all a result of our search for representative artifacts documenting the origins of the space sciences in the United States. Many of them bordered on IGY interests and activities, but did not center on them. Even so, our collection does refl ect the enormous ex- citement and public impact of the fi rst years of the Space Age. We do meet the goal of memorialization, for instance, because at any one time our collections of multiple exam- ples of Explorer 1 and Vanguard 1 spacecraft replicas are on loan to museums across the United States, in Europe, Asia, and Australia. More than a dozen examples of IGY- era artifacts are presently on display at NASM, as well as the new NASM facility at Dulles, the Udvar Hazy Center. So as we ask questions about the IGY during this sea- son of commemoration, at the beginning of the new Inter- national Polar Year 2007? 2008, I hope we will continue to ask how well our collections inform the historical actors, FIGURE 11. Inspection of a replica of Vanguard 1 that had been modifi ed to hold a solar powered audio system for demonstrating the ?beep- ing.? (NASM photograph) 04_DeVorkin_pg035-048_Poles.indd46 4604_DeVorkin_pg035-048_Poles.indd46 46 11/17/08 8:35:51 AM11/17/08 8:35:51 AM PRESERVING THE ORIGINS OF THE SPACE AGE 47 events, episodes and eras that made up the IGY. Where did the expertise come from that allowed our nation to respond to Sputnik and that framed the character of that response? What sorts of technologies were bought to bear? And what was the nature of the infrastructure created to facilitate space-borne research? If our present commemoration ef- forts do not adequately answer these sorts of questions, hopefully future historians will succeed in the effort, asking questions stimulated in part by our material legacy. LITERATURE CITED U.S. House of Representatives. 1946. 70th Congress, 2nd Session, 12 August, Chapter 955, Public Law 79? 722. Department of Space History Collections Rationale, 2005. [The word space was added in the 1960s and serves as basis for the Museum?s Mission Statement, 29 July 1996.] Agreement. 1967. ?Agreement Between the National Aeronautics and Space Administration and the Smithsonian Institution Concern- ing the Custody and Management of NASA Historical Artifacts,? signed 10 March 1967. In the introduction to Space History Divi- sion Collections Rationale, 2005. Bennett, J., 2004. ?Scientifi c Instruments.? In Research Methods Guide, Department of History and Philosophy of Science, 12 March. Cam- bridge, U.K. Cambridge University. Bulkeley, R. P., 1991. The Sputniks Crisis and Early United States Space Policy. Bloomington: Indiana University Press. DeVorkin, D. H. 1992. Science with a Vengeance: How the Military Created the U.S. Space Sciences after World War II. New York: Springer-Verlag. (Reprinted 1993, paperback study edition.) ???. 1996. ?Where Did X-Ray Astronomy Come From?? Rittenhouse, 10: 33? 42. ???. 2006a. ?The Art of Curation: Collection, Exhibition, and Scholarship.? In Showcasing Space, ed. M. Collins and D. Millard, pp. 159? 168. East Lansing: Michigan State University Press. ???. 2006b. ?Space Artifacts: Are They Historical Evidence?? In Critical Issues in the History of Spacefl ight, ed. S. J. Dick and R. D. Launius, NASA SP-2006? 4702, pp. 573? 600. Washington, D.C.: National Aeronautics and Space Administration (NASA). Green, C. M., and M. Lomask, 1970. Vanguard: A History. NASA SP- 4202, Appendix 3, pp. 290? 293. Washington, D.C.: NASA. Hanle, Paul, and Von del Chamberlain, eds., 1981. Space Science Comes of Age. Washington, D.C: Smithsonian Institution. Hirsh, R. 1983. Glimpsing an Invisible Universe: The Emergence of X-ray Astronomy. Cambridge, U.K.: Cambridge University Press. Kennicott, Philip. 2007. ?At Smithsonian, Gay Rights Is Out of the Closet, into the Attic.? Washington Post (8 September 2007), C1; C8. Kopytoff, Igor, 1986. ?The Cultural Biography of Things: Commoditi- zation as Process.? In The Social Life of Things: Commodities in Cultural Perspective, ed. Arjun Appadurai, pp. 64? 91. Cambridge, U.K.: Cambridge University Press. Korsmo, Fae L. 2007. ?The Genesis of the International Geophysical Year,? Physics Today, 70 (2007), 38? 44. ???. ?The Policy Process and the International Geophysical Year, 1957? 1958.? In Smithsonian at the Poles: Contributions to Inter- national Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 23? 34. Washington, D.C.: Smithsonian Institution Schol- arly Press. Ludwig, George H. 1959. ?The Instrumentation in Earth Satellite 1958 Gamma.? MSc diss., February, SUI-59? 3, State University of Iowa, Iowa City. ???. 1960. ?The Development of a Corpuscular Radiation Experi- ment for an Earth Satellite.? Ph.D. diss., August, SUI-60? 12, State University of Iowa, Iowa City. ???. 2005. Letter to D. H. DeVorkin, 26 September. Curatorial Artifact Files A20060086000. Deed of Gift 1476 (26 January 2006). Smithsonian Institution, Washington D.C. Mack, Pamela E., and D. H. DeVorkin. 1982. Conference Report on Pro-Seminar in Space History. Technology and Culture, 23, No. 2(April): 202? 206. Marson, Frank M., and Janet R. Terner. 1963. United States IGY Bibli- ography, 1953? 1960; an Annotated Bibliography of United States Contributions to the IGY and IGC (1957? 1959). Washington, D.C.: National Academy of Sciences-National Research Council. McDougall, Walter A. 1985. The Heavens and the Earth: A Political His- tory of the Space Age. New York: Basic Books. Needell, Allan A. 2000. Science, Cold War and the American State: Lloyd V. Berkner and the Balance of Professional Ideals. Amster- dam: Harwood Academic in association with the National Air and Space Museum, Smithsonian Institution. Pisano, Dominick A., and Cathleen S. Lewis, eds. 1988. Air and Space History: An Annotated Bibliography. New York: Garland. Space Astronomy Oral History Project (SAOHP). 1981? 1986. Smithson- ian Institution, National Air and Space Museum, Archives Division, MRC 322, Washington, D.C. Schorzman, T. A., ed. 1993. A Practical Introduction to Videohistory. Malabar, Fla.: Krieger. Sullivan, Walter A. 1961. Assault on the Unknown; the International Geophysical Year. New York: McGraw-Hill. Van Allen, J. A. 1983. Origins of Magnetospheric Physics. Washington, D.C.: Smithsonian Institution Press. Wilson, J. Tuzo, 1961. ?Foreword by Lloyd V. Berkner.? In I.G.Y., the Year of the New Moons. New York: Knopf. 04_DeVorkin_pg035-048_Poles.indd47 4704_DeVorkin_pg035-048_Poles.indd47 47 11/17/08 8:35:54 AM11/17/08 8:35:54 AM 04_DeVorkin_pg035-048_Poles.indd48 4804_DeVorkin_pg035-048_Poles.indd48 48 11/17/08 8:35:55 AM11/17/08 8:35:55 AM ABSTRACT. The author discusses three archaeological investigations of historic sites in the polar regions. The fi rst site is that of the Solomon A. Andr?e expedition camp on White Island, Svalbard. This fateful ballooning expedition to the North Pole in 1897 was the fi rst experiment in polar aeronautics. Andr?e and his colleagues gave their lives but opened the door to polar fl ight, the backbone of polar logistics today. The other site, East Base, on Stonington Island off the Antarctic Peninsula, served the1939? 1941 U.S. Antarc- tic Service Expedition, under Admiral Richard Byrd, the fi rst U.S. government? sponsored scientifi c and aerial mapping effort in Antarctica. In 1992, a team of archaeologists docu- mented and secured the site that had been recently recognized as an historic monument by the Antarctic treaty nations. The third site is Marble Point on Victoria Land across from Ross Island and McMurdo Station. In conjunction with the IGY 1957? 1958, a massive effort was put into laying out a 10,000-foot year-round runway and creating a fresh water reservoir and other base facilities. It was one of the premier locations for strategic avia- tion in Antarctica. The site was archaeologically surveyed and original engineering docu- mentation from 1956? 1957 offers superb baselines for studying permafrost, erosion, and human disturbances in the Antarctic environment. These types of sites are in situ monu- ments to human courage, ingenuity, and perseverance on a par with NASA?s exploration of space. They require careful management and protection following the same principles as historic sites within the United States and in other nations. INTRODUCTION There is a seemingly limitless public interest in polar exploration. This fas- cination is one of the greatest resources we have for support of polar science: Public enthusiasm for polar history and exploration should be actively acknowl- edged in projects of these kinds. With volunteer help led by professional archae- ologists on regular tour ships, site documentation and cleanup efforts could be carried out on a scale that would otherwise be impossible to achieve. This paper is about international polar history and heritage along with sci- ence and technology in context. We are now embarked on the Fourth Interna- tional Polar Year. In addition to projects in the natural and physical sciences, focusing especially on issues of global change, there are a number of themes in anthropology and archaeology, and new historic archaeology projects are being initiated at both poles. Noel D. Broadbent, Arctic Studies Center, Department of Anthropology, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 112, Washington, DC 20013- 7012, USA (broadbentn@si.edu). Accepted 29 May 2008. From Ballooning in the Arctic to 10,000-Foot Runways in Antarctica: Lessons from Historic Archaeology Noel D. Broadbent 05_Broadbent_pg049-060_Poles.ind49 4905_Broadbent_pg049-060_Poles.ind49 49 11/17/08 8:39:59 AM11/17/08 8:39:59 AM 50 SMITHSONIAN AT THE POLES / BROADBENT The three archaeological studies presented here span 60 years of aviation history from before the period of fi xed-wing aircraft to the International Geophysical Year in 1957? 1958. They also relate to competing national interests in the polar regions including Nordic rivalries, U.S. and German confl icts over Antarctic territories dur- ing World War II, and Cold War strategic thought in the southern hemisphere. Historic archaeology is based on the theories and methods of traditional archaeology but applies these to historic periods (South, 1977; Orser, 2004; Hall and Silliman, 2006). One great advantage of historic archaeol- ogy over prehistoric archaeology is that it can be matched with written sources that may reveal, among other things, the motives behind various endeavors and the details of planning and consequent successes and failures. However, written history tends also to focus on the larger picture of given events and can be biased by political, social, or other concerns of the times. Physical evidence and archaeologi- cal analysis can be used not only to test hypotheses about historical events but also, of equal importance, to provide information about the daily lives of individuals that rarely make it into historical accounts. Global warming, one of the great scientifi c concerns of today regarding polar ecosystems, is having profound impacts on archaeological sites. These sites are subject to increasingly severe weathering and erosion damage. Warmer conditions have also made them more accessible to tourism, as well as to looting. Documentation of these sites is urgently needed. This is exemplifi ed by the Andr?e site in Svalbard, Norway. While such efforts have intrinsic value in and of themselves, the importance of these in- vestigations extends beyond that, inasmuch as the sites themselves can provide baselines for measuring the effects of climate change over time. This is the case of the two Antarctic sites, East Base and Marble Point. THREE CASE STUDIES The fi rst study was an investigation of the S. A. Andr?e ballooning expedition to the North Pole in 1897 (Andr?e, Strindberg, and Fraenkel, 1930). This study was under- taken in 1997? 2000 as a purely scientifi c investigation into the causes of death of these Swedish polar explorers and to document their campsite (Broadbent, 1998; 2000a; 2000b). The centennial of this expedition was 1997, and the project was supported by the Royal Swedish Acad- emy of Science and the Nordic Research Council for the Humanities within the framework of a larger history of science program (Wr?kberg, 1999). The second study was undertaken in 1992 as an Ant- arctic environmental cleanup and cultural heritage man- agement project. The United States Congress allocated environmental funds in 1991 to facilitate the cleanup of old American bases in Antarctica. One of these bases, East Base on the Antarctic Peninsula dating to 1940, is the old- est remaining American research station in Antarctica. The archaeological project was conducted through the auspices of the Division of Polar Programs at the National Science Foundation in collaboration with the National Park Ser- vice (Broadbent, 1992; Broadbent and Rose, 2002; Spude and Spude, 1993). The third study, Marble Point, was directly related to the International Geophysical Year of 1957? 1958. Marble Point is situated across the Antarctic continent from East Base and near the large American research station at Mc- Murdo Sound. An enormous engineering effort had been put into building a 10,000-foot runway at Marble Point. This base was, nevertheless, abandoned when the U.S. Navy found its sea approach extremely diffi cult for supply ships. With the aid of volunteers from McMurdo in 1994, it was possible to map the remains of the SeaBee (Navy Construction Battalion Unit or CBU) base camp at Marble Point (Broadbent, 1994). It was in this context that the science that had gone into the prospecting of the airfi eld became apparent, as well as the impacts that construction had left on the landscape. The combination of history, archaeology, and natural science is an exceptionally productive approach to under- standing the past and applying this knowledge to modern research questions. THE ANDR?E NORTH POLE EXPEDITION IN 1897 The Andr?e expedition is one of the most intriguing in polar history. Solomon August Andr?e (1856? 1897) worked at the Swedish patent offi ce. He had become fasci- nated by ballooning after visiting the U.S. centennial cele- bration in Philadelphia in 1876. On returning to Sweden, he devoted himself to designing hydrogen balloons that could be used for exploration and aerial mapping. His greatest effort was put into the design of the Eagle, which was in- tended to take him and two colleagues to the North Pole using southerly winds. A balloon building and hydrogen generating facility were built on Dane Island in northwest- 05_Broadbent_pg049-060_Poles.ind50 5005_Broadbent_pg049-060_Poles.ind50 50 11/17/08 8:40:00 AM11/17/08 8:40:00 AM FROM BALLOONING TO 10,000-FOOT RUNWAYS 51 ern Svalbard in 1896 (Capelotti, 1999). The fi rst attempt on the North Pole was aborted because of poor winds and gas leakage. A new team consisting of Andr?e, Nils Strind- berg, and Knut Fraenkel (replacing Nils Ekholm) launched in the Eagle on 11 July 1897. They were never seen alive again. Interestingly enough, Nils Ekholm, who quit the project which he felt was too risky, was seen by many as a coward and his decision is still being debated in Sweden. He went on to become one of Sweden?s most prominent meteorologists. In 1930, the bodies of Andr?e, Strindberg, and Fraen- kel were found at their campsite on White Island in eastern Svalbard (Lithberg, 1930). The bodies were returned to Sweden, cremated, and buried in Stockholm following an almost royal funeral and national day of mourning (Lund- str?m, 1997). An exhibit of their equipment was shown at Liljevalch?s Art Hall in Stockholm in 1931 ( Fynden p? Vit?n, 1931). Remarkably, their diaries and notes, as well as undeveloped roles of Kodak fi lm, had survived. A num- ber of negatives could be developed and these images pro- vide a haunting picture of their ordeal. The balloon had gone down on the sea ice on 14 July 1897. Because of icing the balloon could simply not stay aloft and after 33 hours of misery the decision was made to land. They had reached 82? 56H11032 north latitude (Figure 1). They were well equipped with sleds and a boat and from 14 July until 5 October they trekked on the sea ice, fi rst east- wards, making little real distance because of the ice drift, then southwards. They were planning on wintering in a comfortable snow house near White Island and celebrated the king?s birthday on September 18 with a meal of seal meat, liver, kidneys, brains, and port wine. They were in fi ne spirits. Then disaster struck when the ice fractured un- der their snow house. Luckily the weather was good. They systematically gathered their equipment and supplies and were forced to go ashore on White Island where they set up a new camp. The last note in their diaries has a date of 17 October. There is little in the diaries about being in a critical situation. There were no last words or explanations, letters to their loved ones or to their sponsors. It seems that they had no inkling that they were in danger of dying. By comparison, Robert Falcon Scott and his companions took the time to leave a number of letters when they understood their situation. They knew at some future date their camp and bodies would be found. As Andr?e and his companions were now on dry land one would assume that, if they had known that all was lost, they would have done the same. They were well aware that the world had been watching them and considerable national prestige was at stake, par- ticularly since the Swedish king, Oskar II, and Alfred Nobel were offi cial sponsors. Strindberg?s body was found in 1930, half buried in a rock crevice near the campsite, indicating that he had been the fi rst to die. Fraenkel lay dead in the tent and Andr?e lay on the rock shelf above the tent. There have been many theories but no conclusive evi- dence for how or why they had died. The campsite that had been mapped during the recovery in 1930 had been largely buried in snow and ice (Lithberg, 1930). Warming trends in the Arctic made it likely that the site would be better exposed today. A new investigation in connection with the centennial of the expedition was organized and subsequently funded. FIELDWORK IN 1998 The fi rst project expedition to Svalbard was under- taken in 1998 on the Norwegian research vessel Lance through the auspices of the Swedish Polar Research Sec- retariat and the Royal Academy of Sciences. The par- ticipants included science historian Sverker S?rlin, the director of the Andr?e Museum, Sven Lundstr?m, Nor- dic Museum ethnographer and polar historian, Rolf Kjellstr?m, archaeology student Berit Andersson, and the author. The team visited White Island for one day and FIGURE 1. The Eagle shortly after landing on the sea ice at 82? 56? N on 14 July 1897. Andr?e and Fraenkel are looking at the balloon. The image was developed from a roll of Kodak fi lm recovered from White Island in 1930. (Photo by Nils Strindberg, courtesy of Andr?e Museum. Gr?nna) 05_Broadbent_pg049-060_Poles.ind51 5105_Broadbent_pg049-060_Poles.ind51 51 11/17/08 8:40:00 AM11/17/08 8:40:00 AM 52 SMITHSONIAN AT THE POLES / BROADBENT mapped the immediate camp area using standard map- ping techniques and photography. The site, as hoped, was found to be totally free of ice and snow cover and even the margins of the tent were still discernable in the sandy soil. Embedded in the tent depres- sion were frozen remnants of clothing. Driftwood, bam- boo, silk shreds, metal fragments, and other small debris were scattered on the soft sand. Even fragments of bone, certainly polar bear and seal bone, but possibly even hu- man, were found (Figure 2). The area surrounding the site also produced artifacts, including opened food tins from the expedition. With this archaeological potential, a follow-up project was proposed in order to map the site using digital technol- ogy and to conduct a soil chemistry analysis in order to map the non-visible areas of the camp and evidence of camp use. It was reasoned that low levels of phosphates, by-products of defecation, urination, and animal carcasses, could also refl ect length of site use. Magnetic susceptibility would tell us about burning on the site. A pile of driftwood still lies near the tent site and had been collected by the Andr?e team as construction material as well as for fuel. From the initial inspection it was possible to verify the archaeological potential of the site and its surroundings. Although most objects had been collected in the 1930s, the ground was still littered with small debris including silk and bamboo from the balloon and boat, tin openers and various metal fragments. This micro-deposition refl ected the extent of the former campsite. The tent and its half frozen fl oor deposits presented an excellent opportunity for study. Among the most interesting conclusions of the survey was that this campsite had been well chosen. It was on well- drained sandy soil and protected by a rocky outcrop. There was a small stream nearby. In addition, the camp was situ- ated well back from the shore and away from areas where polar bears would prowl. Considerable effort had clearly been put into choosing this spot and substantial piles of logs had also been assembled for dwelling construction and/or fuel. We know from a note in Andr?e?s journal that the camp had been named ?Camp Minna? after his mother. FIELDWORK IN 2000 Having received permission by the Norwegian au- thorities to conduct the study, we returned on the Swedish chartered ship Origo to White Island on 19 August 2000 (Broadbent and Olofsson, 2001). The goal of the investigation of the Andr?e site was to archaeologically document the camp and its surround- ings and to assess the potential causes of their deaths. The men did not appear to be either ill-equipped or desperate on going ashore on 5 October 1897. They had established a good campsite on dry ground. They had fuel for their stoves, ammunition, food, and medicines. With this goal in mind, a new potential for analysis at the site emerged in 2000. Mark Personne, MD, of the Stockholm Poison Center, had just published an article in the Swedish Journal of Medicine in which he assessed the symptoms of the three men that could be gleaned from their diaries (Personne, 2000). Suicide, murder, depres- sion, trichinosis, hypothermia, carbon monoxide poison- ing, alcohol/fuel poisoning, and so on had been proposed in the past, but Personne deduced that the most probable cause of death for all three men was botulism poisoning. Botulism Type E is common in the Arctic and is found on the skin of marine mammals that pick it up from bot- tom sediments and in the near-shore environment. It is tasteless and odorless and among the most deadly poi- sons in nature. Once ingested, death occurs within 24? 36 hours (Personne, 2000, plus references). As a neurotoxin, it affects the central nervous system. For our purposes, al- though the bacteria would be long gone, it was technically possible to identify the botulism toxins in the soils of the camp, namely in the area of the tent fl oor. Testing this theory thus became a new target of the investigation. The fi rst setback was to discover that the campsite was now buried under as much as 30 centimeters of ice. Snows from the previous two winters had melted into a hard mass which could only be hacked out with diffi culty. That evening a heavy snow storm with gale-force winds hit the island and the team was unable to go ashore for two days. Some 20? 30 inches of snow now covered the island and the initial survey plans, which had already rendered several interesting fi nds, including one of the three sled yokes from the expedition, were not possible. The planned excavation of the tent fl oor to test Personne?s theory was also no longer possible. The focus returned to the topo- graphic mapping of the site and sampling for phosphates and magnetic susceptibility. Ground level could still be de- termined with our measuring rod and soil samples could still be collected. WHAT THE SOILS REVEALED The Andr?e camp on White Island is situated at 80? 05H11032 N and 31? 26H11032 E. The camp was about 175m from the shore in sandy terrain and adjacent to a 3m high stone out- crop. The site area covers about 250 square meters. From 05_Broadbent_pg049-060_Poles.ind52 5205_Broadbent_pg049-060_Poles.ind52 52 11/17/08 8:40:02 AM11/17/08 8:40:02 AM FROM BALLOONING TO 10,000-FOOT RUNWAYS 53 the perspective of soil chemistry, the site is analogous to the environment of Late Post Glacial hunters living near the ice margins of Scandinavia 9,000 years ago. In an environ- ment like this, almost all organic and phosphate-containing materials were brought to the site by man or beast. Indi- cations of burning were certainly an indication of human presence. Small groups of hunters slaughtering and subsist- ing on animals left relatively high phosphate deposits in the same types of soils. These inorganic phosphates bind with the soil and remain for thousands of years. Citric acids re- leases the phosphate and this can be measured using colo- rimetric (phosphorous-molybdate)? based methods. Johan Olofsson provided the expertise for soil sampling and anal- ysis (Broadbent and Olofsson, 2001). The human body processes 1? 2 g of phosphorus per day (Devlin, 1986). In addition to the deposition of food residues, especially bones, urination and defecation contrib- ute to the buildup of phosphate in the soil. The longer a site is occupied, the greater will be the phosphate enrichment. This buildup is expected to be greatest in the site center or adjacent to the center. Since this is a relative measure, dif- ferences of 5 percent or more are considered signifi cant as related to offsite normal background sample values. The team collected 240 soil samples in the camp area. Phosphate is measured in phosphate degrees, mg P2O5 per 100 g dry soil. The mean phosphate enrichment in the site center was 17H110069 P? with a range of 38? 2 P?. The highest values were actually to the north of the camp margin and associated with water pooling that was a natu- ral process. The average off site values were 11H1100611 P? with a range of 38? 1 P?. Although the site areas averaged slightly higher than the control samples, this was due to natural drainage on the site rather than human activity. In fact, the map of phosphate values shows that the immediate tent area no enrichment whatsoever. This is strong evidence that they had not been active on the site for very long before death overcame them. Magnetic susceptibility (MS) on the campsite aver- aged 5H110064 SI which is comparable to the off site control measurement of 4H110062 SI. One sample on the site measured 40 SI because of a rusty fl ake in the soil sample and this raised the average campsite value. MS is measured in SI units per 10 g of soil using a Bartington MS2B measure cell (Thomson and Oldfi eld, 1986). Magnetic susceptibility rendered no evidence of fi res and the low phosphate values suggest that the site had hardly been used. The three men had arrived on the island on 5 October 1897 in relatively good shape, searched out an excellent campsite, collected heavy driftwood logs for construction of a hut, and then died within hours of each other. They managed a simple burial of the youngest member of the expedition, Nils Strindberg, and then barely made it back to the tent where they collapsed. Andr?e was found with a Primus Stove, which was easily re-lit after 33 years in the snow. Fraenkel was lying on or beside, not in, his sleeping bag. The Personne hypothesis on botulism poisoning could not be tested but remains the most probable cause of the sudden death of all three men. The lack of fi nal words and letters in the otherwise preserved papers supports the idea as well. With exposure to a neurotoxin, the men would have been quickly immobilized and were apparently un- able to write. Oddly enough, Fraenkel was found still wearing his dark glasses in spite of the low light in early October. Light sensitivity is another symptom of botulism (Personne, 2000). A follow-up sampling of the site might one day help prove this theory. Andr?e and his companions still capture the imagina- tion of the public, especially in Sweden. The fact remains that Andr?e was the fi rst pioneer of polar fl ight and aerial photography in the polar regions. While in some quarters he is viewed as a ?balloonatic? who knew this was going to end in disaster, to others he is revered as a genius of balloon design. In 2000, the same year of the investigation of his campsite on White Island, an English adventurer, David Hempelman Adams, launched his hot air balloon, FIGURE 2. The Andr?e campsite in 1998 was completely exposed. Fragments of the balloon, clothing, and other items were visible. (Photo by Noel Broadbent) 05_Broadbent_pg049-060_Poles.ind53 5305_Broadbent_pg049-060_Poles.ind53 53 11/17/08 8:40:02 AM11/17/08 8:40:02 AM 54 SMITHSONIAN AT THE POLES / BROADBENT the Britannic Challenger, from Longearbyn in Svalbard. He reached the North Pole and most of the way back in 133 hours, thus proving that Andr?e?s plan had been fea- sible all along. Every year the Andr?e campsite is visited by numerous tour groups. We noted during our fi rst visit in 1998 that the soft soils of the site were badly disturbed and the area between the site and the shore was dimpled with thou- sands of deep footprints. EAST BASE AND THE UNITED STATES ANTARCTIC SERVICE EXPEDITION (1939? 1941) In 1939, President Franklin Roosevelt appointed Rear Admiral Richard E. Byrd to command the U.S. Ant- arctic Service Expedition. Roosevelt was well aware of German territorial interests in Antarctica; that same year the German Antarctic Expedition on the seaplane tender Swabenland claimed 200,000 square miles of territory, including Crown Princess Maerta Land, which had been a Norwegian claim. Most important coastal points were marked with swastikas and fl ags dropped from aircraft. Germany invaded Norway in 1940 (Broadbent and Rose, 2002). Two American bases were quickly established in Ant- arctica in 1940: West Base under the command of Paul Siple, and East Base under the command of Richard Black (Black, 1946a, 1946b). West Base was built on the Bay of Whales and has been lost in the sea, but East Base, built on Stonington Island off the Antarctic Peninsula (68? 28H11032 S, 67? 17H11032 W), still stands. East Base was hastily abandoned on 22 March 1941 under the looming threat of war. The 26 men, with a smug- gled puppy and a pet bird? Giant Petrel, later donated the FIGURE 3. The abandoned East Base on Stonington Island, Antarctica. The Science Building with its weather tower and the Bunkhouse build- ing were intact. The crate containing a spare engine for the Curtis Wright Condor biplane can be seen in front of the Science Building. (Photo by Noel Broadbent, 1992) 05_Broadbent_pg049-060_Poles.ind54 5405_Broadbent_pg049-060_Poles.ind54 54 11/17/08 8:40:03 AM11/17/08 8:40:03 AM FROM BALLOONING TO 10,000-FOOT RUNWAYS 55 National Zoo in Washington, D.C.? abandoned the base and fl ew out to the ice edge in their Curtis Wright Condor biplane, which also had to be abandoned. The USS Bear then took them back to the United States, and some nine months later the United States was at war with Japan and Germany. Stonington Island was reoccupied by the Falklands Is- lands Dependencies in 1946 and subsequently used by the British Antarctic Service until 1970 (Walton, 1955). East Base itself, although badly vandalized by ship crews from Chile and Argentina, was reoccupied by the Ronne Ant- arctic Research Expedition (RARE) in 1947? 1948 (Ronne 1949; 1979). The RARE expedition was a private venture under the leadership of Finn Ronne who had been second in command at East Base in 1940. The RARE expedition was unique because this was the fi rst time women (Jackie Ronne and the chief pilot?s wife, Jennie Darlington) were to winter over in Antarctica (Ronne, 1950; Darlington and McIlvaine, 1956). All three expeditions conducted important research and determined that this part of the continent was indeed a peninsula and not an island (Ronne, 1949: English, 1941; Wade, 1945). Interest in the historic value of the site was noted by Lipps in the late 1970s (Lipps, 1976, 1978). East Base was recognized as an historic site (#55) by the Antarctic Treaty Nations in 1989. In 1991, U.S. National Park Service archaeologist, Catherine Blee (later Spude), and NPS historian, Robert Spude, were taken to the island by NSF to conduct a sur- vey and develop a management plan for the site. In 1993, the author led a team, including archaeologist and hazard- ous waste expert, Robert Weaver, and staff from the U.S. Antarctic Research Program Base at Palmer Station and British Antarctic Survey personnel from Rothera Station. We were there to follow through on the NPS recommen- dations. These included cleanup and documentation of debris, removal of hazardous materials, repairs of doors, windows, and roofs and the storage of artifacts. In addi- tion, warning and information signs were made for the site and its buildings and an interpretative panel with a description of the station and historic photographs was put on display in the Science Building (Figure 3). The team worked on the site from 20 February until 3 March 1992. The grounds were cleaned and capped with fresh beach stones, the buildings repaired as far as possible and one building became an artifact storage facility. A collection of some 50 artifacts, including old maps, tools, mittens, fi lms, bottles, bunk plates and dog tags with names, scientifi c specimens, medical supplies and other items were brought back and are now kept at the Naval Historical Center, U.S. Naval Yard, in Wash- ington D.C. (Figure 4). A small museum, one of the most remote museums in the world (Iijima, 1994), was set up in the Science Building with and a brass plaque with the names of the 1940? 1941 and 1947? 1948 expedition members that had been do- nated by the National Geographic Society. A guest book was left in the museum together with an American fl ag. A spare engine for the Curtis Wright Condor biplane still rests in its original crate in front of the museum. There is even a World War I vintage light tank with an air-cooled aircraft engine, a failed experiment in winter traction (Fig- ure 5). The fi rst tourists arrived in 29 December 1993 onboard the Kapitan Khlebnikov. In 1994, Jackie Ronne, widow of Finn Ronne, and their daughter, Karen Tupek, visited the site and left more photos and texts for the mu- seum and the bunkhouse. The East Base project was conducted as a cleanup and historical archaeological project. It was the fi rst U.S. effort in historic archaeology in Antarctica that set a precedent for how sites of these types can be managed (Parfi t, 1993). Sites in the East Base area were made environmentally safe and worthy of tourism and are lasting memorials to polar science. More than 100 cruise ships now visit Antarctica every year and there is still an urgent need to document and manage many historic sites around the continent. Preservation conditions have left items in pristine condi- tion and the continent is littered with aircraft, vehicles, and buildings. This is one of the greatest challenges of cultural resource management and ideally should be con- ducted as collaborative international efforts. FIGURE 4. Noel Broadbent recording a cold-weather mask at East Base in 1992. In the foreground are old weather maps found at the site. (Photo by Michael Parfi t) 05_Broadbent_pg049-060_Poles.ind55 5505_Broadbent_pg049-060_Poles.ind55 55 11/17/08 8:40:06 AM11/17/08 8:40:06 AM 56 SMITHSONIAN AT THE POLES / BROADBENT MARBLE POINT, ANTARCTICA The fi nal project to be discussed is a small archaeo- logical endeavor but one encompassing a huge site, that of a 10,000-foot-long airfi eld and base at Marble Point in Victoria Land, across from Ross Island and the large American base at McMurdo Sound. While making site visits to researchers supported by the social sciences program as NSF, the author had the op- portunity to carry out a survey of Marble Point on 22 Jan- uary 1994. With the help of volunteers from McMurdo, we mapped the site of the Navy Seabee (CBRU) camp es- tablished in 1957. There were numerous foundations of ?Jamesway? huts, trash dumps, oil spills, roads, and other features. Large vehicles, graders, and rollers had also been left behind. A reservoir dam been built at the foot of the glacier (Figures 6 and 7a? e). The U.S. Navy (Operation Deep Freeze I) supported American scientists in Antarc- tica during the 1957? 1958 International Geophysical Year and this base was part of the effort. More than 40 nations participated in the IGY and the Antarctic was studied by teams from the United States, Great Britain, France, Nor- way, Chile, Argentina, Japan, and the USSR. Marble Point was both a research site and an ideal place for an airfi eld. It was also a jumping-off point for researchers working in the Dry Valleys. Rear Admiral George J. Dufek championed the airfi eld construction. In an Airfi eld Feasibility Study fi lm (CNO- 5? 1958), he pointed out the strategic signifi cance such an airfi eld for the southern hemisphere. Aircraft could FIGURE 5. The gap between Stonington Island and the Northeast glacier, 1992. As late as in the 1970s, an ice bridge connected the island to the Antarctic Peninsula; it had been the major reason for choosing the island as a base in 1940. The changes refl ect rapid warming in the region. (Photo by Noel Broadbent) 05_Broadbent_pg049-060_Poles.ind56 5605_Broadbent_pg049-060_Poles.ind56 56 11/17/08 8:40:08 AM11/17/08 8:40:08 AM FROM BALLOONING TO 10,000-FOOT RUNWAYS 57 FIGURE 6. Archaeological map of the Navy SeaBee camp ?North Base? from 1957 at Marble Point, Victoria Land, Antarctica. Map shows locations of ?Jamesway? huts, trash dumps, roadways, and oil-spill areas where vehicles had been parked. (Produced by Noel Broadbent, Janu- ary 22, 1994) 05_Broadbent_pg049-060_Poles.ind57 5705_Broadbent_pg049-060_Poles.ind57 57 11/17/08 8:40:10 AM11/17/08 8:40:10 AM 58 SMITHSONIAN AT THE POLES / BROADBENT FIGURES 7a? e. Selection of items found at Marble Point and the remarkable preservation of paper and fabrics. (a) The newspaper cartoons from 1960 still have bright colors. These simple things, large and small, tell us about daily life and work at the base. (Photo by Noel Broadbent). (a) (b) (c) (d) (e) 05_Broadbent_pg049-060_Poles.ind58 5805_Broadbent_pg049-060_Poles.ind58 58 11/17/08 8:40:14 AM11/17/08 8:40:14 AM FROM BALLOONING TO 10,000-FOOT RUNWAYS 59 be rapidly deployed to South America, Africa, and other points north. The sea ice runways used at McMurdo were limited to only part of the year and had to be rebuilt each season. Further, while clearly advantageous for aviation, the icy coast of Victoria Land proved diffi cult for ships to use and the shores were shallow. The 10,000-foot runway was never fi nished. Notably, however, the fi rst ?wheels on dirt? landing of an aircraft in Antarctica took place here when a Navy VXE-6 squadron Otter landed with Sir Edmund Hillary onboard. He had just reached the South Pole in an overland tractor convoy. Marble Point is a weather station and helicopter re- fueling station today. In the 1980s, when the Chinese program was considering the site as a potential research station, the VXE-6 commander, Captain Brian Shumaker, had new stakes placed along the runway to assert continu- ing American presence there. It remains an American site today, and is, without question, the best place for a year- round airfi eld on the continent. It is a rocky promontory located 50 miles from McMurdo Station, and an adjacent land strip serves as a helicopter refueling station in sup- port of U.S. Antarctic program research. Our survey of the Marble Point area revealed that an enormous engineering study had been conducted there in 1956? 1957 by the U.S. Navy. There were test trenches in the permafrost, test pads established with the sensors still in place, and detailed reports by engineers and scientists. In all, the studies encompassed geology, pedology, perma- frost studies, seismic studies, hydrology, glaciology, sea ice and sea bed studies, and polar engineering. This material provides an unparalleled 50-year baseline for studying changes in the Antarctic environment and the effects of human impacts over time. SUMMARY AND CONCLUSIONS Archaeology offers unique opportunities for research on polar history. In addition to adding new substance to material culture remains and a greater focus on individu- als, the documentation of in situ features is a necessary prerequisite for historic site preservation. The loss of sites in the Arctic due to climate change is an enormous prob- lem and increased erosion along Arctic beaches and riv- ers is destroying thousands of years of Arctic prehistory and history. These regions have been little documented as compared with the lower latitudes. New highly perishable artifacts and human remains are melting out of glaciers around the world. The Andr?e expedition study was conducted as a re- search project on the causes of the deaths of these explor- ers. As an immensely popular fi gure in Nordic history, hundreds of visitors visit the campsite every year, made even more accessible by global warming. Management of sites like this will require greater protection, but at the same time we must help facilitate visitors? needs through marked pathways, signs, and better information. The second project, at East Base, was a cleanup and management effort that also served to document and pro- tect the site. This kind of project should continue to be international in scope. In 1992, John Splettstoesser pub- lished an article in Nature regarding the melting of the Northeast glacier at East Base as evidence of rapid global warming (Splettstoesser, 1992). One of the principal rea- sons Stonington Island has been chosen as a base in 1940 was that the glacier connected the island to the peninsula. Today the glacial bridge has vanished and the island is separated by a wide gap of open water. The bridge had been there as late as the 1970s. Human presence at places like Stonington Island has, in other words, inadvertently given us baseline observations for documenting rapid cli- mate change effects. Finally, Marble Point, even as a small mapping effort, further reveals the indirect scientifi c value of former bases. This site is literally a climate-change data goldmine. The engineering baseline data from sites like this, collected for entirely unrelated reasons, are of great value for under- standing human impacts in polar environments. Polar fl ight plays a major part in these three investiga- tions. Andr?e was the fi rst pioneer of polar fl ight. Byrd was at the forefront of polar aviation and in 1946? 1947, after World War II was also instrumental in Operation High- jump. Ronne mapped 450,000 square miles of Antarctica in 1947? 1948. Marble Point was the site of the fi rst dirt runway landing and still remains the largest land airfi eld ever conceived of in Antarctica. Aviation was at the core of mid-twentieth-century exploration and the transition from dog sleds to aircraft has left an amazing legacy of technology at both poles. The ?old politics? of polar research, and the techno- logical efforts put into them, are closely akin to those of international climate change research of today. The Inter- national Geophysical Year in 1957? 1958 was, after all, conducted during the heat of Cold War. The Arctic Ocean has, once again, drawn the close at- tention of the eight polar nations? the United States, Can- ada, Russia, Sweden, Norway, Finland, Iceland, Denmark/ Greenland? as well as all other nations highly dependent 05_Broadbent_pg049-060_Poles.ind59 5905_Broadbent_pg049-060_Poles.ind59 59 11/17/08 8:40:18 AM11/17/08 8:40:18 AM 60 SMITHSONIAN AT THE POLES / BROADBENT on gas and oil. The Antarctic, as international territory, is still a place where national presence is deemed critical. It would not be an exaggeration to state that the past is truly prelude to the most signifi cant issues of the modern era. To ignore this history is to fail to recognize why the polar regions have long played such an important part in the economic and political history of the western world. EPILOGUE Since the logistical costs of archaeological projects in the polar regions are so great, they are rarely possible to carry out. A potential model for achieving a more comprehensive approach to documentation and cleanup could be to use volunteers under expert supervision on regularly scheduled tourist vessels. This would provide meaningful experiences for the public, facilitate polar heritage management, and offer unique opportunities for the tourist industry. This idea is currently being discussed by a consortium of polar histo- rians and archaeologists. The Smithsonian has several sites of interest that would be ideal places to begin. One is the old town of Barrow, site of U.S. efforts in the original IPY in 1881? 1883, and a second site is Fort Conger on Ellesmere Island, used in 1881 by the Greely Expedition LITERATURE CITED Andr?e, S. A., Nils Strindberg, and Knut Fraenkel. 1930. Med ?rnen mot polen. S?llskapet f?r antropologi och geografi . Stockholm: Albert Bonniers F?rlag. Black, Richard B. 1946a. Narrative of East Base, U.S. Antarctic Expe- dition, 1939? 1941, 27 Dec. 1960, Accession III-NNG-57, R 126, National Archives-CP. Washington, D.C. ???. 1946b. ?Geographical Operations from East Base, United States Antarctic Service Expedition 1939? 1941,? Proceedings of the American Philosophical Society, 89: 4? 12. Broadbent, Noel D. 1992. Project East Base: Preserving Research History in Antarctica. NSF Directions, 5: 1? 2. ???. 1994. An Archaeological Survey of Marble Point, Antarctica. Antarctic Journal of the United States, 29: 3? 6. ???. 1998. Report on a preliminary archaeological investigation of the Andr?e expedition camp on White Island, Spitsbergen, 1998. ?Lance? Expedition of the Swedish Program for Social Science Research in the Polar Regions, 16? 24 August 1998. Manuscript, Department of Archaeology, University of Ume?, Sweden. ???. 2000a. Archaeological Fieldwork at Andr?en?set, Vit?n, Spits- bergen? A Preliminary Report. Swedartic 2000, pp. 62? 64. Stock- holm: Swedish Polar Research Secretariat. ???. 2000b. Expedition Vit?n i Andr?es sp?r. Popul?r arkeologi nr. 4, 2000: 19? 22. Broadbent, Noel D., and Johan Olofsson. 2001. Archaeological Investi- gations of the S. A. Andr?e Site, White Island, Svalbard 1998 and 2000. UMARK 23. Arkeologisk rapport. Institutionen f?r arke- ologi och samiska studier. Ume? universitet, Sweden. Broadbent, Noel D., and Lisle, Rose. 2002. Historical Archaeology and the Byrd Legacy. The United States Antarctic Service Expedition, 1939? 31. The Virginia Magazine of History and Biography 2002, 110(2):237? 258. Capelotti, P. J. 1999. Virgohamna and the Archaeology of Failure. The Centennial of S. A. Andr?e?s North Pole Expedition, ed. U. Wr?k- berg, pp. 30? 43. Stockholm: Kungl. Vetenskapsakademien. Darlington, Jennie, and J. McIlvaine. 1956. My Antarctic Honeymoon: A Year at the Bottom of the World. New York: Doubleday and Company. Devlin, T. M. ed. 1986. Textbook of Biochemistry with Chemical Cor- relations. Second edition. New York: Wiley. English, R. A. J. 1941. Preliminary Account of the United States Antarc- tic Expedition, 1939? 1941. Geographical Review, 31: 466? 478. Fynden p? Vit?n. Minnesutst?llning ?ver S. A. Andr?e, Nils Strindberg och Knut Fraenkel anordnad i Liljevalchs konsthall. 1931. Liljev- alchs konsthall, katalog nr. 90. Stockholm. Hall, M., and S. Silliman, eds. 2006. Historical Archaeology. Oxford, U.K.: Blackwell. Iijima, G. C. 1994. Our Most Remote Museum. Odyssey, 3: 41? 42. Lipps, J. H. 1976. The United States, ?East Base,? Antarctic Peninsula. Antarctic Journal of the United States, 11: 215. ???. 1978. East Base, Stonington Island, Antarctic Peninsula. Antarc- tic Journal of the United States, 1978: 231? 232. Lithberg, Nils. 1930. ?The Campsite on White Island and Its Equip- ment.? In Med ?rnan mot polen, ed. S. A. Andr?e, Nils Strindberg, and Knut Fraenkel, pp. 189? 227. Stockholm: S?llskapet f?r antro- pologi och geografi . Lundstr?m, Nils. 1997. ?Var position ?r ej synnerligen god . . . Andr?e expeditionen i svart och vitt.? Stockholm: Carlssons Bokf?rlag. Orser, Charles E. 2004. Historical Archaeology. Upper Saddle River, N.J.: Pearson Education. Parfi t, Michael, and Robb Kendrick. 1993. Reclaiming a Lost Antarctic Base. National Geographic Magazine, 183: 110? 26. Personne, Mark. 2000. Andr?e-expeditionens m?n dog troligen av botu- lism. L?kartidningen 97, 12: 1427? 1431. Ronne, Finn. 1949. Antarctic Conquest: The Story of the Ronne Expedi- tion 1946? 1948. New York: G. P. Putnam?s Sons. ???. 1950. Women in the Antarctic, or the Human Side of Scientifi c Expedition. Appalachia, 28: 1? 15. ???. 1979. Antarctica, My Destiny: A Personal History by the Last of the Great Polar Explorers. New York: Hastings House. South, Stanley. 1977. Method and Theory in Historical Archaeology. New York: Academic Press. Splettstoesser, John. 1992. Antarctic Global Warming? Nature, 355: 503. Spude, Catherine Holder, and Robert L. Spude. 1993. East Base Historic Monument, Stonington Island/Antarctic Peninsula: Part I: A Guide for Management; Part II: Description of the Cultural Resources and Recommendation, NPS D-187, National Park Service. Denver: United States Department of the Interior. Thomson, R., and F. Oldfi eld. 1986. Environmental Magnetism. Lon- don: Allen and Unwin. Wade, F. A. 1945. An Introduction to the Symposium on Scientifi c Re- sults of the United States Antarctic Service Expedition, 1939? 1941. Proceedings of the American Philosophical Society, 89: 1? 3. Walton, E. W. K. 1955. Two Years in the Antarctic. London: Lutterworth Publishers. Wr?kberg, Urban, ed. 1999. The Centennial of S. A. Andr?e?s North Pole Expedition. Stockholm: Royal Swedish Academy of Sciences. 05_Broadbent_pg049-060_Poles.ind60 6005_Broadbent_pg049-060_Poles.ind60 60 11/17/08 8:40:19 AM11/17/08 8:40:19 AM ABSTRACT. The founding of the Smithsonian in 1846 offered the promise of scientifi c discovery and popular education to a young country with a rapidly expanding western horizon. With its natural history and native cultures virtually unknown, Smithsonian Re- gents chartered a plan to investigate the most exciting questions posed by an unexplored continent at the dawn of the Darwinian era. Prominent issues included the origins and history of its aboriginal peoples, and this thirst for knowledge that led the young institu- tion into America?s subarctic and Arctic regions. The Yukon, Northwest Territories, and Alaska were among the fi rst targets of Smithsonian cultural studies, and northern regions have continued to occupy a central place in the Institution?s work for more than 150 years. Beginning with Robert Kennicott?s explorations in 1858, Smithsonian scientists played a major role in advancing knowledge of North American Arctic and Subarctic peoples and interpreting their cultures. Several of these early enterprises, like the explo- rations, collecting, and research of Edward Nelson, Lucien Turner, John Murdoch, and Patrick Ray in Alaska and Lucien Turner in Ungava, either led to or were part of the fi rst International Polar Year of 1882? 1883. Early Smithsonian expeditions established a pattern of collaborative work with native communities that became a hallmark of the institution?s northern programs. This paper presents highlights of 150 years of Smith- sonian work on northern peoples with special attention to themes that contributed to Smithsonian Arctic studies during International Polar/Geophysical Year events, especially 1882? 1883 and 2007? 2008. HISTORICAL CONTRIBUTIONS The International Polar Year (IPY) 2007? 2008 provides an opportunity to explore how the Smithsonian has served for the past 150 years as a repository of Arctic knowledge and a center for northern research and education. When Robert Kennicott arrived in the Mackenzie District in 1859 to make natural history and ethnology collections for the Smithsonian, science in the North American Arctic was in its infancy. By the time the fi rst IPY began in 1882, the Smithsonian had investigated parts of the Canadian Arctic and Subarctic and the Mackenzie District. Further, it had sent naturalists to the Northwest Coast, the Aleutians, western Alaska, and nearby Chukotka in Siberia and was on its way toward developing the largest well- documented Arctic anthropologi- cal and natural history collection in the world. By its close in 1883? 1884 major William W. Fitzhugh, Arctic Studies Center, Department of Anthropology, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 112, Washington, DC 20013- 7012, USA (Fitzhugh@si.edu). Accepted 9 May 2008. ?Of No Ordinary Importance?: Reversing Polarities in Smithsonian Arctic Studies William W. Fitzhugh 06_Fitzhugh_pg061-078_Poles.indd61 6106_Fitzhugh_pg061-078_Poles.indd61 61 11/17/08 8:40:39 AM11/17/08 8:40:39 AM 62 SMITHSONIAN AT THE POLES / FITZHUGH IPY- related fi eld programs in Barrow, Ellesmere Island, and Ungava had been or were nearly completed and col- lecting projects in Kodiak and Bristol Bay were underway. The cumulative results established the Smithsonian as the pre- eminent scholarly institution of its day in the fi elds of northern natural and anthropological science. In the 125 years since IPY- 1, northern collecting, re- search, publication, and education programs have given the Smithsonian an Arctic heritage of immense value to scholars, Natives and northern residents, and interested public around the world. It is a world, moreover, in which Arctic issues have steadily moved from the exoticized pe- riphery of global attention to a well- publicized central focus, as a result of changes in geopolitics, climate, and governance. We are hearing about the north now more than ever before, and this trend is accelerating as global warming strikes deeper into polar regions, transforming oceans, lands, and lives. Elsewhere I explored how nineteenth- century Smith- sonian Arctic scientists laid the foundation for the fi eld of museum anthropology (Fitzhugh, 1988a; 2002a) and pub- lic presentation and exhibition of Arctic cultures at the In- stitution during the past 125 years (Fitzhugh, 1997). Here I review some of the themes that contributed to Smithson- ian Arctic studies during International Polar/Geophysical Year events, especially 1882? 1983 and 2007? 2008. My purpose is not only to illustrate how long- term Smithso- nian research, collecting, and exhibition has contributed to Arctic social and natural science, but also to explore how these historical assets contribute to understanding a region undergoing rapid, dynamic social and environmen- tal change. FOUNDATIONS OF ARCTIC SCIENCE Heather Ewing?s recent book, The Lost World of James Smithson (Ewing, 2007), reveals the Smithsonian?s reclusive founder as more intellectual and politically ac- tive than previously thought but provides few clues as to what his bequest mandate intended. Accordingly, setting the course for the young Smithsonian fell to its fi rst Sec- retary, Joseph Henry (1797? 1878), and his scientifi c as- sistant, Spencer Baird (1823? 1887), who followed Henry as Secretary and presided during the years of IPY- 1. Both Henry and Baird shared in establishing natural science and cultural studies at the Smithsonian and gave early prior- ity to northern studies, which Henry judged ?of no ordi- nary importance? (Smithsonian Institution Annual Report [SIAR], 1860:66). In fact, during the Smithsonian?s earli- est years, it is surprising how much energy went into re- search and publication on Arctic subjects, including Elisha Kent Kane?s meteorological, tidal, and magnetic studies; McClintock?s and Kane?s searches for Franklin; and solar observations and natural history collecting in Labrador and Hudson Bay during the 1850s. Drawing on his previous experience as a regent of the New York University and his association with ethnologists Henry Schoolcraft and Lewis Henry Morgan, Henry was instrumental in laying groundwork for what was to be- come the fi eld of museum anthropology at the Smithson- ian (Fitzhugh, 2002a). In fact, Henry believed that cultural studies would eventually develop into a discipline as rigor- ous as the natural sciences, and for this reason instructed Baird to include ethnology among the tasks of natural- ists he hired as fi eld observers and collectors. Baird be- lieved taxonomic and distributional studies of animals and plants in northwestern North America would reveal rela- tionships with Asia and lead to understanding their origins and development, and he came to believe that ?ethnologi- cal? collections could also reveal deep history. In 1859, a gifted young prot?g?e of Baird?s named Robert Kennicott (1835? 1866) became the fi rst of ?Baird?s missionaries? (Rivinus and Youssef, 1992:83; Fitzhugh, 2002a) sent north to begin this grand task (Lindsay, 1993). Kennicott spent 1859 to 1862 in the Hudson Bay Territory and Mackenzie District making the fi rst carefully documented natural history collections from any North American Arctic region. Assisted by Natives and Hudson Bay Company agents, he also collected more than 500 eth- nological specimens from Inuvialuit (Mackenzie Eskimo) and Dene Indians, as swell as linguistic data, myths and oral history, and ethnological observations. In his report to the Regents on 1861, Henry noted the collections being submitted by the factors of the Hudson?s Bay Company, ?taken into connexion with what Mr. Kennicott is doing, bid fair to make the Arctic natural history and physical geography of America as well known as that of the United States? (SIAR, 1862:60). A year later Henry reported (SIAR, 1863:39), ?This enterprise has terminated very favorably, the explorer having returned richly laden with specimens, after making a series of observations on the physical geography, ethnology, and the habits of animals of the regions visited, which cannot fail to furnish materi- als of much interest to science.? In 1860, when Kennicott fi rst began to explore west from the Mackenzie into Russian America territory, only southern and western Alaska had been previously ex- plored ethnologically by Russians, and northern and east- ern Alaska was nearly unknown (Sherwood, 1965; James, 06_Fitzhugh_pg061-078_Poles.indd62 6206_Fitzhugh_pg061-078_Poles.indd62 62 11/17/08 8:40:39 AM11/17/08 8:40:39 AM ?OF NO ORDINARY IMPORTANCE? 63 1942). Ethnological collecting had been conducted spo- radically in coastal and southwest Alaska since ca. 1800 by Russia and its agents [e.g., Lavrentii Zagoskin (1842? 1844)]. In 1839? 1849, purposive but not scientifi cally di- rected ethnological museum collecting was carried out by Ilya G. Voznesenskii for the Russian Academy of Sciences (Black, 1988; Fitzhugh, 1988a; Kuzmina, 1994; Fitzhugh and Chaussonnet, 1994). In 1865, the Western Union Telegraph Company was pushing an overland telegraph line to Europe via the Yu- kon River, Bering Strait, and Siberia. That year Baird asked Kennicott to direct the scientifi c activity of the survey with assistance from William Healy Dall, Henry W. Elliott, and several other naturalists (Figure 1; Collins, 1946; Fitzhugh and Selig, 1981). While making the fi rst scientifi cally doc- umented American collections from interior and coastal Alaska north of the Aleutians, the project collapsed after Kennicott?s death on the Yukon River in May 1866, and the subsequent completion of a transatlantic cable by a rival company the following July. Nevertheless, the West- ern Union survey produced the fi rst scientifi cally docu- mented American collections from Alaska, trained the fi rst American scholars of Alaska, and led to the fi rst English- language books on ?the great land? written by Henry W. Elliott (1886), Frederick Whymper (1869), and William Healy Dall (1870). In the early 1870s, Baird and Dall began to implement a more ambitious Alaskan collecting venture. Realizing the Smithsonian could not fi nance a sustained endeavor by itself, Baird recruited government agencies like the U.S. Army Signal Service, Hydrographic Offi ce, and War De- partment to employ Smithsonian naturalists as weather and tidal observers at government stations throughout the newly purchased territory. Their activities produced a vast collection of meteorological, geographical, natural histori- cal, and anthropological data that, supplemented by pho- tography after 1880 (Fitzhugh, 1998c), laid the scientifi c bedrock for later studies in Alaska and northern Quebec. FIGURE 1. Frederick Whymper?s illustration of the Western Union Telegraph Survey team battling ice on the Yukon River below Nulato in 1866 in Eskimo umiaks in spring 1866. (From Whymper, 1869) 06_Fitzhugh_pg061-078_Poles.indd63 6306_Fitzhugh_pg061-078_Poles.indd63 63 11/17/08 8:40:40 AM11/17/08 8:40:40 AM 64 SMITHSONIAN AT THE POLES / FITZHUGH While Baird?s collecting and publication programs were an unqualifi ed success, only part of Baird?s and Henry?s original plan was ever realized. Documentation of endangered cultures and languages fulfi lled their science plan and provided materials for further study, following Henry?s expectation that new methods and techniques would lead to creating a ?hard? science of anthropology. This prospective view from the mid- nineteenth century is still our optimistic view at the turn of the twenty- fi rst century but it probably will never be realized. Progress in anthropological science has come in different directions: Ethnology and cultural anthropology have not merged with the natural or hard sciences as Baird and Henry pre- dicted. Rather, Smithsonian anthropology proceeded to develop in other directions: the study of human remains and forensics; archaeology (long one of Henry?s interests but one that was impossible to conduct with the method and theory available in the late nineteenth century); and another set of anthropological fi elds not imagined by them at all? heritage, ethnicity, and cultural identity. SMITHSONIAN ACTIVITIES IN IPY- 1 (1881? 1884) As the fi rst IPY approached, the Smithsonian had conducted work throughout most regions of Alaska south of the Bering Strait: James G. Swan had collected on the Northwest Coast in the 1850? 1880s; Robert Kennicott in British America in 1859? 1862 and in interior Alaska, 1865? 1866; William Healy Dall in western Alaska and the Aleutians, 1865? 1885; Lucien Turner in St. Michael, 1871? 1877, and the Aleutians, 1877? 1878; and Edward W. Nelson had just completed studies in the Yukon, Kus- kokwim, and Bering Strait in 1877? 1881 (Figure 2). Sev- eral other collecting projects were proceeding in southern Alaska, but none had been carried out north of Bering Strait. The 1881 voyage of the Revenue Cutter Corwin briefl y visited the coast between the Bering Strait and Bar- row with a scientifi c team including Edward W. Nelson, John Muir, and Irving Rosse, reaching as far east as the Inuit settlement of Ooglamie at today?s Barrow, and as far west as Herald and Wrangel Islands in the Chukchi Sea and Wankarem on the Arctic coast of Siberia. The visit to Barrow was only two days, and Nelson?s diary notes that he had diffi culty making collections and gathering infor- mation. Barrow people had been dealing with European whalers for almost three decades and knew how to drive hard bargains. As documented in accompanying papers in this vol- ume, the Smithsonian?s major collection focus in IPY- 1 were Barrow and Ungava Bay. Given the Institution?s interest in Alaska, Barrow became the primary target of a major effort directed by Lt. Patrick Henry Ray with the assistance of John Murdoch, a Smithsonian employee who carried out ethnological studies (Ray, 1885; Murdoch, 1892; Burch, 2009, this volume; Crowell, 2009, this volume). The Barrow project fi lled the last major gap in the Smithsonian?s sur- vey coverage of Alaska and, second only to Nelson?s work, made the most important contribution to science. Living for two full years in a weather station near the native village, Murdoch concentrated his efforts on ethnological collecting and reporting. Like Nelson, he collected vocabularies and linguistic data, but he did not venture out on long trips with Native guides or show much interest in oral history and mythology (Burch, 2009, this volume; A. Crowell, Arctic Studies Center, personal communication, 2007). Murdoch?s more remote approach led him to be duped by Natives who sold him artifacts they constructed hastily for sale or had composited from unmatched materials, inserting stone scrapers in ivory handles that had no blades or embellishing ancient ivory objects with new designs. Nevertheless, while Murdoch was not as perceptive a cultural observer, his work received great notice in the developing profession of anthropology. Boas? The Central Eskimo (1888a), also published by the Smithsonian, made Murdoch?s Ethnological Results of the Point Barrow Ex- pedition (1892) the fi rst study of a Western Arctic Eskimo FIGURE 2. Ceremonial mask representing Tunghak, Spirit of the Game, collected by Edward W. Nelson from the Lower Yukon River, undergoing conservation in 2003. (NMNH 33118) 06_Fitzhugh_pg061-078_Poles.indd64 6406_Fitzhugh_pg061-078_Poles.indd64 64 11/17/08 8:40:48 AM11/17/08 8:40:48 AM ?OF NO ORDINARY IMPORTANCE? 65 group. More importantly, Murdoch?s monograph directly addressed the scholarly debates about Eskimo origins and migrations and utilized a scientifi c method, making spe- cifi c comparisons of Alaskan Eskimo customs and mate- rial objects with those known from the Canadian Arctic and Greenland. When Nelson?s (1899) monograph, The Eskimos About Bering Strait, appeared several years later, it was not fully appreciated by anthropologists because Nelson was a biologist and his highly descriptive study did not address the Eskimo origin controversy (Fitzhugh, 1988b). As a result, until the 1980s most scholars did not comprehend the importance of differences between Yup?ik and I?upiat material culture, art, and language that these monographs clearly illustrated. Even Boas, who made much of cultural continuities in Raven mythology, art, and folklore between Northwest Coast and Northeast Asia (Boas, 1903, 1905, 1933; Bogoras, 1902), failed to recog- nize that these features were also present in the geographi- cally intermediate Yup?ik Bering Sea and I?upiat Eskimo area; he continued throughout his life to promote the idea that Eskimos were recent arrivals to the Bering Strait from Canada (Boas, 1888b, 1905; see below). It was not until much later that Yup?ik Eskimo culture began to be under- stood as a distinct Eskimo tradition with stronger ties to the south than to I?upiat and other Arctic coast Eskimo cultures, and with a legacy from ancient Eskimo cultures of the Bering Sea (Fitzhugh, 1988c; Dumond, 2003). Other projects also made important contributions to the Smithsonian?s IPY-1 program even though they were not part of the offi cial IPY agenda. Charles MacKay?s col- lections from the Signal Service Station at Nushugak in Bristol Bay (1881? 1883) contained fascinating materials from the border between Yup?ik, Aleut, and Alutiiq cul- tures, complementing S. Applegate?s materials from Un- alaska and nearby regions (1881? 1885). However, none of these collections was ever published or exhibited, and the early ethnography of this boundary region is still poorly known today. More prominent is the work of Wil- liam J. Fisher who contributed materials from southern Alaska and Kodiak Island throughout the 1880s (Crowell, 1992). Like Applegate and MacKay, Fisher did not pre- pare reports; however, his collection became the subject of intensive recent study and exhibition (Crowell et al., 2001). Two other projects also were ?offi cial? Smithson- ian IPY ventures (Krupnik, 2009, this volume). Adolphus Greely?s ill- fated scientifi c explorations at Fort Conger, Lady Franklin Bay, Ellesmere Island in High Arctic Can- ada, were a massive undertaking organized by the U.S. Sig- nal Service in 1882? 1883 (Barr, 1985), for which Spencer Baird served as scientifi c advisor. Although the expedition ended in disaster, it obtained important scientifi c observa- tions. Many of the weather, aurora, and meteorological observations were archived at the Smithsonian, which also received a few natural history and ethnological specimens, along with the team?s scientifi c instruments. Today the Bar- row and Fort Conger weather records serve as important benchmarks for long- term study of climate change (Wood and Overland, 2006). The third offi cial Smithsonian IPY- 1 fi eld study, Lucien Turner?s ethnological work among the Innu and Inuit of Ungava Bay in northern Quebec during 1882? 1884, produced a trove of important ethnological materials from both Innu (Naskapi) and Inuit (Eastern Es- kimo) groups, as well as natural history and photographic records (Turner 1894; Loring, 2001a, 2009, this volume). BUILDING ON IPY-1: MUSEUM EXHIBITION, RESEARCH, AND ANTHROPOLOGICAL THEORY By 1890, the great era of synoptic Smithsonian natural history- based collecting had passed and attention began to be devoted to publishing, collection work, building a spe- cialized scientifi c staff, and presenting American cultures to the world. Following the 1893 Chicago Columbian Expo- sition, a series of world?s fairs exhibited living Eskimos and other northern peoples together with displays of museum collections to wide audiences in Chicago, New Orleans, St. Louis, Buffalo, and other locations. The Smithsonian?s Arctic ethnography collections were featured in many of these exhibitions, and some of the displays were later in- stalled in the Smithsonian?s permanent galleries. This was the era of the dramatic life- group diorama reconstructions pioneered by the Smithsonian?s famous artist- geologist William Henry Holmes. His Polar Eskimo group for the Buffalo fair in 1901 became one of the most popular exhib- its after the National Museum of Natural History opened in 1910 and remained on view for nearly 100 years (Figure 3; Ewers, 1959:513? 525; Fitzhugh, 1997). Concurrent with growth of exhibitions and new ar- chitecture on the Washington Mall, the Smithsonian began to build its curatorial staff and hire its fi rst professionally trained anthropologists. The Bureau of American Ethnol- ogy founded by John Wesley Powell in 1879 as a center for anthropological fi eld surveys, research, and publication (Hinsley, 1981) was the fi rst Smithsonian entity staffed by anthropologists. In 1891, Powell?s linguistic surveys and subsequent synthesis produced the fi rst linguistic map of North America, a tour de force of early museum research that seemed to establish language as the guiding structure 06_Fitzhugh_pg061-078_Poles.indd65 6506_Fitzhugh_pg061-078_Poles.indd65 65 11/17/08 8:40:51 AM11/17/08 8:40:51 AM 66 SMITHSONIAN AT THE POLES / FITZHUGH for cultural diversity. However, by 1893, Otis Mason?s study of material culture collections across North America showed both congruence and discontinuity across linguistic boundaries (Figure 4). Eskimo collections fi gured promi- nently in his work, demonstrating gradual stylistic changes in dress, implements, kayaks, and other equipment (Ma- son, 1891, 1896, 1902; Ewers, 1959:513? 525)? except in Greenland, where 200 years of exposure to European culture had produced a radical departure from traditional Central and Western Eskimo clothing and design. Mason concluded that tribal material culture and language groups were not always synchronous, as Powell had supposed, but more closely followed C. Hart Merriam?s biogeographic life zones. But even here discontinuities resulted from external infl uence, migration, language capture and loss, and other cultural and historical factors. Mason?s ?culture area? con- cept was a direct outgrowth of museum- based research on the Smithsonian?s Arctic IPY- 1 collections and remains one of the underpinnings of anthropological theory today. As analysis of the Smithsonian?s pan- North American Arctic collections progressed, it seemed that anthropology was drifting further from the pure science Henry predicted it would become. NEW SCIENCE ARRIVES: PHYSICAL ANTHROPOLOGY AND ARCHAEOLOGY These gropings toward the development of anthropo- logical science did not become fully professionalized until Ales wedgesubscript Hrdlic wedgesubscript ka (1869? 1943), the father of American physi- cal anthropology, joined the Smithsonian in 1903 and be- FIGURE 3. The Polar Eskimo of Smith Sound, Greenland, display? created by William Henry Holmes for the Buffalo Exposition in 1901 and on exhibit since then at the National Museum of Natural History? was dismantled in October 2004. Robert Peary Jr. of Qaanak (at right), the Inuit grandson of the American explorer who collected some of the materials for this exhibit, was present for the event. (Courtesy Department of Anthropology, Smithsonian Institution) 06_Fitzhugh_pg061-078_Poles.indd66 6606_Fitzhugh_pg061-078_Poles.indd66 66 11/17/08 8:40:51 AM11/17/08 8:40:51 AM ?OF NO ORDINARY IMPORTANCE? 67 gan to study the question of Indian and Eskimo origins with new methods and discipline applied to human skel- etal remains. Hrdlic wedgesubscript ka?s Alaskan work utilized methods of fi eld collecting offended native people, and are seen to- day as outrageously insensitive, and his scientifi c results, while interesting in their day, have been largely superseded (Scott, 1994). Perhaps his most lasting contribution was recruitment of T. Dale Stewart and Henry B. Collins to the Smithsonian staff in the mid 1920s. Stewart refi ned Hrdlic wedgesubscript ka?s osteological methods and inherited Hrdlic wedgesubscript ka?s mantle while Collins took a different path, bringing ar- chaeology into the forefront of studies of Eskimo ori- gins through his pioneering stratigraphic excavations on St. Lawrence Island in the 1930s (Collins, 1937, 1951). Here an unbroken sequence of changing artifact forms, art styles, house types, and economies demonstrated a long history of local development interrupted periodically by Asian infl uences over the past 2,000 years. Hrdlic wedgesubscript ka?s, Stewart?s, and Collins? early work in Alaska was con- ducted largely without reference to the Smithsonian?s early IPY collections and research products and without reference to IPY- 2 and IGY 1957? 1958 program efforts, with which Smithsonian scientists had little involvement (Krupnik, 2009, this volume). PUBLIC ?DISCOVERY? OF ESKIMO ART In 1973, the Institution?s IPY- 1 ethnological collec- tions from Barrow and Collin?s prehistoric archaeologi- cal materials from Bering Strait resurfaced suddenly and dramatically with the refi ned cachet of ?Eskimo art? when FIGURE 4. This spear thrower display was assembled by Otis Mason to demonstrate spatial changes in artifact types across culture and space, analogous to biological species distribution. North American Eskimo throwers (bottom row) are arranged from Alaska (left) to Greenland and Labrador (right). These and many other Eskimo artifacts demonstrate systematic style change from west to east. (Courtesy Department of Anthropology, Smithsonian Institution) 06_Fitzhugh_pg061-078_Poles.indd67 6706_Fitzhugh_pg061-078_Poles.indd67 67 11/17/08 8:40:54 AM11/17/08 8:40:54 AM 68 SMITHSONIAN AT THE POLES / FITZHUGH the National Gallery of Art opened its groundbreaking ex- hibition The Far North: 2000 Years of American Indian and Eskimo Art (Collins, 1973). C. D. Lewis, curator of sculpture at the Gallery, was assigned to curate the exhi- bition, and I assisted his search for northern art among the Smithsonian?s Arctic collections. The experience was life- changing. Acquaintance with the collections, coupled with the phenomenal success of the exhibit, convinced me that the Smithsonian ?attic? housed treasures of inter- est not only to anthropologists and Native constituencies but also to a far broader audience. To paraphrase former Smithsonian Secretary S. Dillon Ripley?s reference to the Institution?s musical instrument collections, we needed to take the Arctic treasures out of their storage cabinets and make them ?sing.? In the late 1970s, I began to do that, and with William C. Sturtevant started meeting with anthropologists from the Soviet Academy of Science?s Institute of Ethnography to plan an exhibit on the cultures of Siberia and Alaska. Political diffi culties caused periodic delays, and it did not open until 1988, in the early phase of the Gorbachev rev- olution known as perestroika (Fitzhugh, 2003). During the years while Crossroads was gestating, Susan Kaplan and I created an exhibit based on the ethnological col- lections made by Edward W. Nelson in 1877? 1881 from western Alaska and Bering Strait. Inua: Spirit World of the Bering Sea Eskimo (Fitzhugh and Kaplan, 1983) ex- plored the art, culture, and history of the Yup?ik peoples of southwest Alaska. The exhibit (Figure 5) and catalog illustrated the extraordinary beauty and workmanship of Yup?ik and Bering Strait I?upiat culture. After opening at the Smithsonian in 1982, Inua toured to Anchorage, Fair- banks, Juneau, and other cities in North America. Later, Kaplan and I created a mini- Inua version that toured to FIGURE 5. E. W. Nelson collections from 1877? 1881 in a hunting ritual display in Inua: Spirit World of the Bering Sea Eskimo. (1983 photo- graph; courtesy Arctic Studies Center, Smithsonian Institution) 06_Fitzhugh_pg061-078_Poles.indd68 6806_Fitzhugh_pg061-078_Poles.indd68 68 11/17/08 8:40:57 AM11/17/08 8:40:57 AM ?OF NO ORDINARY IMPORTANCE? 69 small museums and culture centers in Alaska, Canada, and Greenland (Fitzhugh and Kaplan, 1983). Eventually a third version, ?Euro- Inua? (Figure 6), was developed by Susan Rowley for a tour across eastern and northern Europe and Iceland (Rowley, 1988). After the long hiatus following Collins? work in the 1930s, these 1980s exhibits and publications brought Smithsonian Alaska collections to a wide audience in North America and to the rest of the world, especially to Alaska residents and native villages. They also brought us to the attention of Ted Stevens, U.S. Senator from Alaska. Early in 1980 while I was preparing Inua, I had occasion to give his wife, Ann Stevens, a tour of the Smithsonian?s Alaskan collections in what was then a very dusty Natural History Museum attic. A few days later, the Senator called for his own tour, during which he remarked, ?Bill, we have to fi nd a way to get these collections back to Alaska.? The IPY- 1 and other early Alaskan collections indeed had a captivating power, and it was growing year by year. That tour and the senator?s remark gave me my marching or- ders for the next twenty- fi ve years and in time led to a dedicated Smithsonian program reconnecting its historic collections with Alaska and its Native peoples. As I explored the Smithsonian attic, I was amazed to discover how little the collections were known. In those days the Smithsonian?s attic was a virtual King Tut?s tomb before excavation? quiet, dusty, and full of splendid things! The Smithsonian had never hired an Arctic ethnol- ogist, and Hrdlic wedgesubscript ka, Collins, and Stewart had not strayed far from their osteological and archaeological disciplines. The few scholars aware of the collections knew them only from small black and white illustrations in Nelson?s and Murdoch?s monographs. Ronald Senungetuk, an art- ist on the staff of the University of Alaska in Fairbanks who came to Washington in 1981 to consult on the Inua exhibit, may have been the fi rst Alaska Native to inspect them fi rsthand. During the late 1980s, we completed arrangements to launch Crossroads of Continents: Cultures of Siberia and Alaska (Fitzhugh and Crowell, 1988). The exhibit (Fig- ure 7) was based on a reciprocal exchange that paralleled the history of the collections: The earliest objects from Alaska had been gathered during the Russian? America era and had been stored at the Museum of Anthropology and Ethnography in Leningrad since the 1840s, whereas the earliest Siberian materials had been gathered by Franz Boas? Jesup North Pacifi c Expedition and were held by the American Museum of Natural History in New York. FIGURE 6. Catalogs issued for the mini- Inua exhibitions that toured small museums in Alaska in 1983? 1984 (Fitzhugh and Kaplan, 1983) and in Europe 1988? 1989 (Rowley, 1988). (2008 photo- graph; courtesy Arctic Studies Center, Smithsonian Institution) FIGURE 7. Crossroads of Continents combined Russian collections from Alaska with American collections from Siberia in an integrated exhibition featuring the history, culture, and art of the peoples of the North Pacifi c rim. (1988 photograph; courtesy Arctic Studies Cen- ter, Smithsonian Institution) 06_Fitzhugh_pg061-078_Poles.indd69 6906_Fitzhugh_pg061-078_Poles.indd69 69 11/17/08 8:41:00 AM11/17/08 8:41:00 AM 70 SMITHSONIAN AT THE POLES / FITZHUGH Logistics, geography, political barriers, and lack of pub- lication and scholarly exchange, to say nothing of native awareness, had made it impossible to synthesize a larger view of the traditional cultures of the North Pacifi c ?cross- roads? region. Every one of the 650 specimens was jointly selected and researched by teams from both sides during yearly visits fi nanced by the International Research and Exchanges Board (IREX). Co- curated with Aron Crow- ell and assisted by Val?rie Chaussonnet, Sergei Arutiunov, Sergei Serov, Bill Holm, James VanStone, and many others, including Igor Krupnik, this exhibit put us in direct contact with Soviet scholars and laid the groundwork for future research, publication, conferences, and exhibit ventures. The show toured in Alaska and the lower 48 and was the fi rst joint U.S.? Soviet exhibition in which American and Soviet/Russian materials were published in a single catalog and comingled in a single display (Fitzhugh, 2003). In the early 1990s, following precedent established by the mini- Inua exhibits, Val?rie Chaussonnet organized a small version of Crossroads called Crossroads Alaska that toured towns across Alaska (Chaussonnet, 1995; Carlo et al., 1995). Unfortunately, when it came time to transfer the large Crossroads exhibit to Russia, economic and se- curity conditions had deteriorated so much that the tour was cancelled. Nevertheless, as a substitute, in 1996 we ar- ranged for a smaller exhibit Crossroads Siberia (Krupnik, 1996) to tour cities in the Russian Far East, curated by Igor Krupnik, who had come to work at the Arctic Studies Center in 1991. This was probably the fi rst anthropology exhibit to travel in Siberia, as well as the fi rst Alaska Na- tive artifacts to be seen in the Russian Far East. A WIDENING FOCUS: ARCTIC STUDIES CENTER AND CIRCUMPOLAR ANTHROPOLOGY In 1988, the Smithsonian received congressional sup- port for creating a special unit called the Arctic Studies Center (ASC) in the National Museum of Natural History, enabling a series of new research, education, and publi- cation ventures that have been described in annual ASC newsletters and on the website (www.mnh.si.edu/arctic). Building anew on the IPY- 1 legacy and new Crossroads partnerships, in 1992 a 10- year archival research and pub- lication project titled ?Jesup II? was initiated with United States, Russian, Japanese, and Canadian partners (Krupnik and Fitzhugh, 2001; Kendall and Krupnik, 2003). To ful- fi ll Boas? original vision of the scope of the Jesup expedi- tion (Boas, 1903)? a vision thwarted by bureaucracy and logistics during the expedition and by Soviet fi at excluding the Ainu from the Crossroads exhibition? with Japanese and Ainu scholars, we produced a comprehensive Ainu ex- hibition and catalog drawing on collections and archives in museums in North America and Japan (Fitzhugh and Dubreuil, 1999). Concurrent with the opening of fi eld opportunities in Russia after 1990, the ASC began a re- study of culture themes that motivated circumpolar theories of Arctic peo- ples in the early twentieth century (Gjessing, 1944; Bogoras, 1902, 1929; Fitzhugh, 1975; Dumond, 2003) with new fi eldwork in the lower Ob River and Yamal Peninsula. Sup- ported by a grant from Amoco Eurasia Corporation, we conducted archaeological surveys and ethno- archaeology of Nenets reindeer- herders in Western Siberia (Fedorova et al., 1998; Fitzhugh, 1998a; 1998b; Fitzhugh and Golovnev, 1998; Haakanson, 2000; Fedorova, 2005) combined with several museum- focused heritage projects, exhibits, and catalogs (Krupnik and Narinskaya, 1998; Krupnik, 1998; Pika, 1998). Subsequently, with Andrei Golovnev and Vlad- imir Pitul?ko, we searched for pre- Eskimo sites eastward from Yamal along the Arctic coast to Bering Strait, inspired by Leonid Khlobystin?s pioneering work (Khlobystin, 2005) and assisted excavations at an 8,000- year- old Mesolithic site on Zhokhov Island (Pitul?ko, 2001). Although the re- sults did not reveal evidence of proto- Eskimo culture, they helped explain why earlier researchers believed Eskimo ad- aptations and art had originated in these regions (Larsen and Rainey, 1948; Fitzhugh, 1998a) and helped fi ll a large gap in circumpolar archaeology (Fitzhugh, 2002b). Later our research and public programs gap in the North At- lantic was fi lled by production of a major exhibition titled Vikings: the North Atlantic Saga, which opened in 2000 and toured to various locations in North America (Fitzhugh and Ward, 2000). REVITALIZING THE SMITHSONIAN? ALASKA CONNECTION Beginning in the early 1990s, the reprinting of Nelson?s and Murdoch?s monographs and new interest created by the Alaska tours of Inua and Crossroads and their mini- exhibit versions resulted in a major revitalization of the Smithsonian? Alaska connection. Unlike the earlier focus on collecting and research, these efforts were based on collection interpretation, education, and public access. Whereas earlier work involved primarily a one- way trans- 06_Fitzhugh_pg061-078_Poles.indd70 7006_Fitzhugh_pg061-078_Poles.indd70 70 11/17/08 8:41:05 AM11/17/08 8:41:05 AM ?OF NO ORDINARY IMPORTANCE? 71 fer of Alaskan objects and information to Smithsonian coffers for use in research, publication, and exhibitions, the emerging emphasis used the Smithsonian treasures to engage Native groups and individuals in two- way collab- orative studies and publication, along with joint curation of exhibitions, museum training, and re- documentation of the Smithsonian?s early object and archival collections ( Fienup- Riordan, 1996, 2007; Loring, 1996). The overwhelming interest exhibited by Alaskans to early Smithsonian collections helped spark a revival of in- terest in traditional native culture, and in the early 1990s we began to explore the idea of opening a regional offi ce to formalize a new permanent Smithsonian? Alaska con- nection. In April 1994, we opened an offi ce at the Anchor- age Museum of History and Art and shortly after, Aron Crowell joined the ASC as local director and launched archaeological research, museum training, exhibition, and teaching projects. As the Alaska offi ce took shape, back in Washington the ASC staff collaborated on collection projects with the National Museum of Natural History (NMNH) Repatria- tion Offi ce, which was working with Alaska Native groups on the return of human remains collected by Hrdlic wedgesubscript ka, Stewart, Collins, and others in the early 1920? 1930s. More than 3,500 skeletal remains were transferred back to Native groups between 1990? 2007, together with asso- ciated grave goods and religious objects (Bray and Killion, 1994; http://anthropology.si.edu/repatriation). ASC staff began to work with Native groups to document old ar- chival and ethnographic collections from Alaska, and by 2001 these ?knowledge repatriation? projects (Crowell et al., 2001; Loring, 2001b, 2008; Krupnik, 2004, 2005) blossomed into the Alaska Collection Project, bringing Alaska Natives into contact with Smithsonian collections for intensive study and re- documentation, with the ulti- mate aim of loaning them back to Alaska for study and exhibition. In 1994 Crowell began an exhibition project with the William J. Fisher Alutiiq ethnographic collection from the Kodiak Island area gathered in 1880? 1885 during the fi rst IPY era, but never previously published or exhibited. The resulting exhibit? Looking Both Ways: Heritage and Identity of the Alutiiq People? and its catalog and web- site, co? curated by Crowell with Alutiiq leaders and or- ganized with the Alutiiq Museum (Crowell et al., 2001; Pullar, 2001), helped catalyze the movement by Alutiiq peoples to rejuvenate Kodiak cultural traditions in art, oral history, language, and material culture (Crowell, 2004; Clifford, 2004). The collections and observations garnered as well as the re- publication (with new data) of the Nelson, Murdoch, and Turner monographs, and presentation of exhibitions and new illustrated catalogs have proven important both for science and historical legacies. For instance, without historical baseline information from archaeological sites, it is impossible to determine the signifi cance of climate shifts or assess the effects of long- term cultural and en- vironmental change. Further, preservation of traditional artifacts, customs, and oral histories and presentation of these materials through exhibitions and other venues have given Alaska Natives a window into a past that had been largely forgotten or was considered irrelevant to the mod- ern day (Kaplan and Barsness, 1986; Fitzhugh, 1988c; Chaussonnet, 1995). Now nearing its fi fteenth year, the Smithsonian rela- tionship with the Anchorage Museum is poised to take another giant step forward. With funding from the Ras- muson Foundation and others, the Anchorage Museum has constructed a new wing to house an expanded ASC of- fi ce and research suite. A major part of this wing will be a Smithsonian exhibition hall displaying nearly 650 anthro- pological objects loaned from the Smithsonian?s National Museum of Natural History and the National Museum of the American Indian (Figure 8). The collections have FIGURE 8. Unangan consultants (from left) Daria Dirks, Marie Turnpaugh, Vlaas Shabolin, and Mary Bourdukofsky study a painted wooden shield from Kagamil in the Aleutian Islands, Alaska (ASCN11:3). Native experts? young and old? have helped to re- document the objects with new information, stories, songs, and native language, and the process has helped to fi nd new routes to the past. (2003 photograph; courtesy Arctic Studies Center, Smithsonian Institution) 06_Fitzhugh_pg061-078_Poles.indd71 7106_Fitzhugh_pg061-078_Poles.indd71 71 11/17/08 8:41:06 AM11/17/08 8:41:06 AM 72 SMITHSONIAN AT THE POLES / FITZHUGH been selected by teams of exhibit designers, conservators, and Alaska Native experts under the curatorial direction of Aron Crowell and will open in 2010 (Figure 9). A pilot exhibit titled Sharing Knowledge and a website of the same name have been created (Figure 10; http://alaska . si.edu) and expanded educational programs are planned as part of a new phase of the Smithsonian?s commitment to Alaska and its cultures and peoples. INTO THIN AIR: ARCTIC STUDIES AND IPY 2007? 2008 The convergence of the Smithsonian?s effort to forge a new relationship with Native and other constituencies in Alaska and across the circumpolar north featured in many current International Polar Year projects is hardly a coincidence. Just as the Smithsonian?s work with north- ern peoples has evolved over the past fi fteen years, so too have biologists, oceanographers, and other natural scien- tists begun to recognize the need for active involvement of northern residents in the enterprise of polar science. For the fi rst time since IPY- 1, the 2007? 2008 IPY includes so- cial science as a major research focus as well as? for the FIGURE 10. The Arctic Studies Center?s Alaska offi ce prepared Sharing Knowledge: the Smithsonian Alaska Collections (http:// alaska.si.edu) in collaboration with Native elders and Second Story Interactive Studio provides interactive assess to historic collections enriched by new Native documentation and oral history. FIGURE 9. The Ralph Applebaum Associates rendition of the Native Cultures exhibition to open in 2010 in the Smithsonian Gallery of the expanded Anchorage Museum. 06_Fitzhugh_pg061-078_Poles.indd72 7206_Fitzhugh_pg061-078_Poles.indd72 72 11/17/08 8:41:09 AM11/17/08 8:41:09 AM ?OF NO ORDINARY IMPORTANCE? 73 fi rst time in polar research? direct participation by north- ern peoples (Krupnik et al., 2004; Krupnik and Hovelsrud, 2006; Allison et al., 2007; NOAA, 2008; www.ipy.org). Back in Washington, as preparations for the 2007? 2008 IPY began to take shape amid the growing realiza- tion that climate warming was altering the Arctic world in ways that had never been imagined, the ASC curated an exhibit exploring the forces at work in this regional expression of global change (Figure 11). Arctic: A Friend Acting Strangely (2006), produced by the ASC with as- sistance from NOAA, NSF, NASA climate scholars as a major component of the U.S. Interagency Arctic Research Policy Committee?s SEARCH (Study of Arctic Environ- mental Change) Program, presented the science of Arctic warming and its effects on marine and terrestrial systems, animals, and people. Special attention was given to human observations of changes in the Arctic, such as rising tem- peratures, reductions in permafrost and sea ice, increases in coastal erosion, shorter winters and longer summers, shifts in animal distributions, and the possible local extir- pation of some species important for human subsistence (NOAA, 2008; http://forces.si.edu/arctic; www.arctic .noaa .gov). The exhibit helped focus the national climate debate from exclusive attention to geophysical drivers of global warming to its human and social ?face? by illustrating the massive changes underway in the Arctic, long before such effects are expected to become pronounced at lower lati- tudes. Many of these issues are subjects of ongoing IPY sci- ence initiatives. The ASC activities most closely associated with these efforts are found in Aron Crowell?s research into culture and climate history in the Gulf of Alaska re- gion (Crowell and Mann, 1998; Crowell, 2000; Crowell et al., 2003), William Fitzhugh?s (1998a, 2002b; Fitzhugh and Lamb, 1984) work on long- term culture and environ- mental change and human-environmental interactions in the circumpolar region, and Igor Krupnik?s (Krupnik and FIGURE 11. View of National Museum of Natural History exhibition Arctic: A Friend Acting Strangely, documenting the impacts of climate change on Arctic animals, landscapes, peoples, and cultures in ancient and modern times. (Photo by Chip Clark, NMNH) 06_Fitzhugh_pg061-078_Poles.indd73 7306_Fitzhugh_pg061-078_Poles.indd73 73 11/17/08 8:41:12 AM11/17/08 8:41:12 AM 74 SMITHSONIAN AT THE POLES / FITZHUGH Jolly, 2002; Huntington et al., 2004; Krupnik et al., 2004; Krupnik, 2006) collaborative projects on indigenous obser- vations of sea ice, animal, and climate change in the Ber- ing Sea, and Stephen Loring?s (1996, 1998; 2001a; 2001b; 2008; Loring and Rosenmeier, 2005) work with indigenous community science and education. It is clear that global warming, as dramatically dem- onstrated during the 2007 summer melt season, is going to be the most serious environmental issue facing the world in the coming century. Building upon the Smithsonian?s long history of anthropological and archaeological collect- ing and research, the ASC has the capability for deep- time and broad panoramic studies of culture and environmen- tal change. The Institution?s ethnographic collections and archival records provide information on how northern peoples in many regions of the north have adapted to re- gional variation and changing conditions. Its archaeologi- cal collections, particularly from Alaska? as well as its recent long- term studies in the Eastern Arctic and Subarc- tic, the Russian North, Scandinavia, and most recently in Mongolia? provide cultural and environmental informa- tion on past changes of climate, environment, and culture that form the basis for studies and educational programs informing current conditions and trends. Movements of prehistoric and historic Indian and Eskimo cultures in Lab- rador, responses of past and present sea mammal hunters on St. Lawrence Island to changing sea ice and animal dis- tributions, and cultural changes seen in Russia and Scandi- navia have been taking place for thousands of years. One of the challenges of IPY 2007? 2008 is to apply knowledge of these and similar records to the conditions we are facing today, and to assist local government and people living in these regions in making sensible choices for the future. The Smithsonian?s long history of northern studies from Kennicott?s fi rst steps in the Mackenzie District in 1858 to the modern day; its collections, research, and public programs; and its contemporary collaboration with northern communities and peoples give it unique capac- ity for contributions in this IPY and in this time of rapid social and environmental change. After 150 years of draw- ing upon the north as a source of collections and scholarly research exemplifi ed in the Institution?s fi rst IPY efforts, Smithsonian science and education have shifted the po- larity of its collecting, science, and educational activities back into the north so that its resources can contribute to meeting the challenges that lie ahead through the direct involvement of Alaskan and other Arctic people. Smithso- nian scholars still research and publish at the forefront of their fi elds, curate collections, and work with the public in many capacities; but sharing the Smithsonian?s historic collections and archives and opening its facilities as venues for education and expanded awareness have become the guiding star for this new phase in our history. Nothing could be more important or more worthy of the course established by our founders in the earliest days of the In- stitution, for whom Arctic research is, as Henry deemed, ?of no ordinary importance.? ACKNOWLEDGMENTS This paper owes much to the long- term support and intellectual stimulation provided by the Smithsonian?s dedicated staff, particularly of the Department of Anthro- pology and its Collections Program staff, who have man- aged and maintained the Arctic collections and supported the many exhibitions and outreach programs conducted by the ASC since the early 1980s. Susan Kaplan (Bowdoin College), Val?rie Chaussonnet, Aron Crowell (ASC), Nata- lia Fedorova (Museum- Exhibit Center, Salekhard, Russia), Andrei Golovnev (Institute of History and Archaeology, Ekaterinburg, Russia), Susan Kaplan (Bowdoin College), Igor Krupnik (ASC), Stephen Loring (ASC), Noel Broad- bent (ASC), Vladimir Pitul?ko (Institute of the History of Material Culture, St. Petersburg, Russia), Susan Rowley (University of British Columbia), Ruth Selig (NMNH), Elisabeth Ward (University of California, Berkeley; for- merly with ASC), and Patricia Wolf (Anchorage Museum of History and Art) have been of great assistance and inspiration as friends and colleagues and contributed to programs and ideas expressed above. Of course, nothing would have been possible without the tradition of scholar- ship and service maintained by the Smithsonian since its founding. 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WEBSITE RESOURCES Smithsonian Institution http://anthropology.si.edu http://anthropology.si.edu/repatriation www.mnh.si.edu/arctic International Polar Year www.ipy.org Arctic: A Friend Acting Strangely http://forces.si.edu/arctic National Oceanographic and Atmospheric Administration www.arctic.noaa.gov Sharing Knowledge, ASC Alaska Collection Project http://alaska.si.edu 06_Fitzhugh_pg061-078_Poles.indd77 7706_Fitzhugh_pg061-078_Poles.indd77 77 11/17/08 8:41:18 AM11/17/08 8:41:18 AM 06_Fitzhugh_pg061-078_Poles.indd78 7806_Fitzhugh_pg061-078_Poles.indd78 78 11/17/08 8:41:18 AM11/17/08 8:41:18 AM ABSTRACT. The following pages review four research trips to Smithsonian collections made by Yup?ik community members between 1997 and 2003. In each case, Yup?ik elders had the opportunity to examine ethnographic material gathered from southwest Alaska in the late-nineteenth and early-twentieth centuries. Most had seen similar objects in use locally when they were young and provided rich commentary not only on the sig- nifi cance of particular tools and pieces of clothing, but on the traditional way of life and worldview that fl ourished in southwest Alaska in the 1920s and 1930s. Although much has changed since these elders came of age?the introduction of organized religion, for- mal education, and a wage economy?a rich and vibrant oral tradition remains. Through sharing knowledge in collections as well as working with museum professionals to bring objects home to Alaska for exhibition, elders seek not only to remind their younger generations of their rich heritage, but to declare the ingenuity and compassion of their ancestors to all the world. INTRODUCTION Yup?ik Eskimo men and women fi rst gained awareness of Smithsonian col- lections in 1982, with the opening of Bill Fitzhugh and Susan Kaplan?s ground- breaking exhibition, Inua: Spirit World of the Bering Sea Eskimo (Fitzhugh and Kaplan, 1982) in Anchorage. Prior to the opening of the Yup?ik mask exhibit, Agayuliyararput/Our Way of Making Prayer in 1996, elders worked with pho- tographs of objects, but few entered museums to see the real thing. Since then, Yup?ik elders have had unprecedented opportunities to visit and view Smith- sonian collections, including one- and two-week research trips to the National Museum of the American Indian in 1997 and 2002, and the National Museum of Natural History in 2002 and 2003. Inua and Agayuliyararput opened doors, and those who entered found an unimagined array of artifacts, which most had viewed only briefl y when they Ann Fienup-Riordan, Arctic Studies Center, 9951 Prospect Drive, Anchorage, AK 99516, USA (riordan@alaska.net). Accepted 9 May 2008. Yup?ik Eskimo Contributions to Arctic Research at the Smithsonian Ann Fienup-Riordan 07_Fienup-Riordan_pg079-088_Pole79 7907_Fienup-Riordan_pg079-088_Pole79 79 11/17/08 8:36:21 AM11/17/08 8:36:21 AM 80 SMITHSONIAN AT THE POLES / FIENUP-RIORDAN were young. All were deeply moved by what they saw. Elders also recognized the potential power of museum col- lections to communicate renewed pride and self-respect to a generation of young people woefully ignorant of the skills their ancestors used to survive. Finally, in 2003 the Calista Elders Council began to actively search for ways to bring museum objects home. Repatriation was not the issue, as ownership of objects was not the goal. Rather the Council sought ?visual re- patriation?? the opportunity to show and explain tradi- tional technology to contemporary young people. The re- sults of their work in Smithsonian collections has not only enriched our understanding of nineteenth-century Yup?ik technology in unprecedented ways but also laid the foun- dation for the exhibition Yuungnaqpiallerput (The Way We Genuinely Live): Masterworks of Yup?ik Science and Survival, bringing Yup?ik materials home to Alaska in this Fourth International Polar Year. SOUTHWEST ALASKA The Yukon-Kuskokwim region, a lowland delta the size of Kansas, is the traditional homeland of the Yupiit, or Yup?ik Eskimos. The region?s current population of more than 23,000 (the largest native population in Alaska) lives scattered in 56 villages, ranging between 200 and 1,000 persons each, and the regional center of Bethel with a population of nearly 7,000 (Figure 1). Today this huge re- gion is crosscut by historical and administrative boundar- ies, including two dialect groups, three major Christian denominations, fi ve school districts, two census areas, and three Alaska Native Claims Settlement Act (ANCSA) re- gional corporations. Villages each have an elementary and a secondary school, city government or traditional coun- cil, health clinic, church or churches, airstrip, electricity, and, in some cases, running water. With 14,000 speakers of the Central Yup?ik language, the Yupiit remain among the most traditional Native Americans. The subarctic tundra environment of the Bering Sea coast supports rich fl ora and fauna. An impressive variety of plants and animals appears and disappears as part of an annual cycle of availability on which Yup?ik people focus both thought and deed. Millions of birds nest and breed in the region?s ample wetlands, including geese, ducks, and swans. Annual migrations of salmon and herring are major resources for both riverine and coastal hunters. Halibut, fl ounder, tomcod, whitefi sh, capelin, pike, needlefi sh, smelt, and blackfi sh seasonally appear in coastal waters and tun- dra lakes and sloughs, and seals, walrus, and beluga whales return each spring. Land animals abound, including moose, caribou, bear, fox, otter, Arctic hare, muskrat, and beaver, and edible greens and berries are plentiful during summer months. Prehistorically this abundance supported the de- velopment and spread of Inuit culture, and some scholars have called the coast the ?cradle of Eskimo civilization.? The abundance of plants and animals in southwest Alaska allowed for a more settled life than in other parts of the Arctic. Hundreds of seasonal camps and dozens of winter settlements lined riverine highways that link com- munities to this day. Like the northern Inuit, the coastal Yupiit were nomadic, yet their rich environment allowed them to remain within a relatively fi xed range. Each of at least a dozen regional groups demarcated a largely self- suffi cient area, within which people moved freely through- out the year in their quest for food. Far from seeing their environment as the insentient provider of resources avail- able for the taking, many Yupiit continue to view it as responsive to their own careful action and attention. ELDERS IN MUSEUMS: FIELDWORK TURNED ON ITS HEAD This year, 2007, marks the beginning of the Fourth International Polar Year. September 2007 also saw the opening of the exhibition, Yuungnaqpiallerput/The Way We Genuinely Live: Masterworks of Yup?ik Science and Survival, in Bethel, Alaska. Yuungnaqpiallerput was devel- oped by the Calista Elders Council in collaboration with the Anchorage Museum with funding from the National Science Foundation. It is based on over a decade of work in museums by Yup?ik men and women, including four seminal research trips to Smithsonian collections. The fol- lowing pages share highlights of how and why elders came to the Smithsonian, what they learned, and what they hope to do with this knowledge. Yuungnaqpiallerput (Fienup-Riordan, 2007) stands squarely on the shoulders of the successful partnership that gave rise to the Yup?ik mask exhibit Agayuliyararput/Our Way of Making Prayer in 1996 (Fienup- Riordan, 1996). That exhibit was the culmination of efforts to understand the meaning and power of nineteenth-century masks, in- cluding masks collected by Edward Nelson and Lucien Turner, from the Yup?ik point of view. The cornerstone of that exhibition, as with Yuungnaqpiallerput, was informa- tion eloquently shared by Yup?ik elders during both pri- vate and public conversations, remembering the masked dances they had seen when they were young. Some had seen photographs of masks, but few had entered muse- 07_Fienup-Riordan_pg079-088_Pole80 8007_Fienup-Riordan_pg079-088_Pole80 80 11/17/08 8:36:21 AM11/17/08 8:36:21 AM YUP?IK ESKIMO CONTRIBUTIONS TO ARCTIC RESEARCH 81 ums to see the real thing until after the exhibit opened in Toksook Bay in January 1996. Following Agayuliyararput, Yup?ik men and women have had unprecedented opportunities to visit museums and view collections. The fi rst ?Yup?ik delegation? to do serious work with Smithsonian collections was a group of six elders who, along with myself and Marie Meade, traveled to the Bronx storage facility of the National Mu- seum of the American Indian (NMAI) for two weeks in April 1997. NMAI had invited the elders to New York as thanks for what they had shared during the Yup?ik mask exhibit, which was then on display in New York. Our visit marked the fi rst time Smithsonian staff extended such an invitation to Alaska Native elders. Organized in large part by Mary Jane Lenz, NMAI housed us, fed us, and shared with elders as many objects as they could during the time we had together. Agayuliyararput opened museum doors, and those who entered found an unimagined array of artifacts, including hunting equipment, clothing, and the tools of daily life. Ironically, the objects elders found least interesting were the masks, which most had viewed only briefl y when they were young. Grass socks, stone tools, and fi sh-skin cloth- ing, however, excited enormous interest. All were deeply FIGURE 1. Map of Southwest Alaska, 2008. (Patrick Jankanish and Matt O?Leary) 07_Fienup-Riordan_pg079-088_Pole81 8107_Fienup-Riordan_pg079-088_Pole81 81 11/17/08 8:36:22 AM11/17/08 8:36:22 AM 82 SMITHSONIAN AT THE POLES / FIENUP-RIORDAN moved by what they saw and spoke repeatedly about the skill required to make and use each item. Viewing col- lections for the fi rst time, the late Willie Kamkoff (April 1997) of Kotlik remarked: ?Seeing these things after we arrived, our ancestors were so ingenious in making hunt- ing tools. They didn?t have iron tools, only ciimat [rocks]. Their tools weren?t sharp, but they were amazing.? Elders also recognized the potential power of museum collections to communicate renewed pride and self-respect to a generation of younger Yup?ik men and women woe- fully ignorant of the skills their ancestors used to survive. Refl ecting on his visit to NMAI, Paul John (September 1998) of Toksook Bay said: We saw many objects when we visited the museum in New York, but I couldn?t leave the adze and an ax for a long time. I kept going back to look at them in awe, realizing that they had been used by someone long before metal and nails were intro- duced. The ax had an ivory blade with a wooden handle shiny from constant sweat and oil from the hands that held it. . . . The objects in museums are not insignifi cant. If we live us- ing them as our strength, we will get closer to our ancestors? ways. And when we are gone, our grandchildren will continue to live according to the knowledge they have gained. Elders were deeply engaged by the full range of Yup?ik technology. This point was brought home during the sec- ond major Yup?ik foray into Smithsonian collections, a visit to NMAI?s Cultural Resources Center in August 2000. Three elders had been invited to choose objects for inclusion in the new museum?s planned exhibit, Our Uni- verses. During the fi rst four days of their visit, they sat in a conference room and carefully described their way of life to curator Emil Her Many Horses and staff. On the last day they were invited into storage areas where all 1,000 Yup?ik objects in the museum?s collections were spread be- fore them. The elders were asked to walk through the room and choose the things that they felt best refl ected the tradi- tional Yup?ik view of the world. The Yup?ik group was the last of eight groups to make such selections. Some, like the Pueblo, had chosen 40 pieces, including many sacred ones. Others made smaller selections. The Yup?ik group circled the room, pointed enthusiastically at everything recog- nizably Yup?ik, and chose more than 300 objects. Masks were of interest, but the technology that had allowed their ancestors to survive was of primary importance. This trip, planned with the help of the Calista Elders Council (CEC) and fully funded by NMAI, was preceded by a two-day gathering in Bethel, where fi ve elders an- swered questions and discussed their view of the world with Emil Her Many Horses and Mary Jane Lenz. This meeting was foundational, not only for NMAI staff but for CEC, the region?s primary heritage organization representing the 1,300 Yup?ik elders sixty-fi ve and older. CEC had just be- gun actively documenting traditional knowledge in 1999, both during their annual Elder and Youth Conventions and interviews with individual elder experts. The format NMAI chose? a small group of elders focusing discussion on a specifi c topic? was an inspiration. Supported by NSF, the CEC has since held more than two dozen two- and three- day gatherings on topics chosen by CEC?s board of elders, including ones on relational terms, discipline techniques, migratory waterfowl, and fall survival skills. In my 30 years work in the region, the last fi ve years have been both the most satisfying and the most productive, seeking to answer questions Yup?ik people themselves are posing. Aron Crowell, Director of the Arctic Studies Cen- ter?s Anchorage offi ce, organized the third Yup?ik visit to Smithsonian collections in 2002. There, three elders spent one week examining and commenting on a rich range of objects Aron selected for inclusion in the Arctic Stud- ies Center?s Anchorage exhibition, scheduled to open in 2010. Photographs and elders? observations from this im- portant trip are available on the Arctic Studies Center?s website, Sharing Knowledge. Again, it was deeply moving to see how much exploring collections meant to individ- ual elders. Eighty-year-old John Phillip Sr. from Kongiga- nak wanted to come so badly that when stormy coastal weather closed in, he drove his snow machine more than 100 miles to Bethel to make his fl ight. February 2003 was the last Yup?ik trip to work in the Smithsonian Institution?s Museum Support Center (MSC). Frank Andrew, the single most knowledgeable elder I have every known, had been unable to come to Washington, D.C., in 2002 due to his wife?s death. Members of the CEC staff had worked with Frank at museums in Bethel and Anchorage, but he wanted to see objects specifi cally from the Canineq (lower coastal) area of the Bering Sea. So, we organized this one-week trip just for him. Frank was accompanied by his son, Noah, as well as two knowl- edgeable elders he felt comfortable with. The Smithsonian Community Scholars program funded our travel, with ad- ditional support from the Calista Elders Council. Prior to our week at MSC, I searched records and pre- selected close to 300 objects from Canineq (Nelson, 1899). The MSC staff was generous in supplying records and print outs to make this long-distance selection possible. During our stay, we recorded close to 30 hours of discussion, all in Yup?ik, producing more than 1,000 pages of transcripts. We had an additional adventure when the blizzard of 07_Fienup-Riordan_pg079-088_Pole82 8207_Fienup-Riordan_pg079-088_Pole82 82 11/17/08 8:36:31 AM11/17/08 8:36:31 AM YUP?IK ESKIMO CONTRIBUTIONS TO ARCTIC RESEARCH 83 2002 hit D.C. the night before we were to fl y home. Frank had noted that the Yup?ik name for the small snowfl akes he saw falling was taqailnguut, ones that don?t stop? and they didn?t. Washington shut down and planes didn?t fl y for another three days. The elders were unconcerned, as waiting out storms was a routine part of life. Food was a potential problem, but Noah and I went ?hunting? every afternoon for whatever Pennsylvania Avenue could pro- vide? tins of smoked oysters, roast chickens, canned soup. During the days, we gathered in our largest room and told stories. I kept the recorder running, producing what Marie affectionately referred to as the Blizzard Tapes. Not only have elders traveled to Smithsonian collec- tions but Smithsonian collections have come home also. During our last trip to MSC, NMAI photo archivist Donna Rose gave us a fat binder of copies of the won- derful photographs made by Dr. Leuman M. Waugh, re- cently saved from oblivion by Igor Krupnik and Stephen Loring, among others (Krupnik and Loring, 2002; Fienup- Riordan, 2005). An especially moving account was re- corded while looking at Waugh?s photo of men hunting in kayaks near Frank Andrew?s hometown (Figure 2). I naively asked Frank if he felt pride the fi rst time he used a kayak. Here is what he said. There were two men [Qilkilek and Puyulkuk] who picked on me. One was my grandfather, and the other was my cross- cousin. Getting a kayak didn?t make me feel important. One of them picked on me very hard. He said that I would only eat catches that I got from other people and not my own and that I would only wear clothing that was handed down. The other one asked me why I got a kayak, one that would rot before the blood of my catch soaked it. These things that I heard were not things that make you feel good. That was how I felt when I got a kayak, and I stopped sleep- ing. I didn?t want to be [the way they said I would be]. I learned everything about paddling before I got a kayak by using one that wasn?t mine, and I was taught how to hunt as well. I was fi lled with eagerness and the will to succeed. I got a kayak during summer. They put it up on stilts. I would even go and check on it at night when it was windy, being afraid it might blow away because that old man bothered me so much. . . . Then during [the following] spring they began to go down to the ocean. I stopped sleeping, wondering when he would let me go. They eventually began to catch sea mammals. After that, my late older sister told me that they were going to take me. I was so ecstatic because I loved to be by the ocean when I went to fetch their catches. And we would be reluctant to go back up to land sometimes. It was always calm. We were getting ready to go down. . . . We walked, pull- ing our kayak sleds at night, and we got to the ocean just at daybreak. I saw the person who ridiculed me. His name was Qilkilek, and he was in a kayak. I never forgot what he said to me. When he saw me he said, ?What are you here for? Go back up and be a urine-bucket dumper. You won?t catch anything.? I didn?t answer. Frank then left to hunt with his two cousins. Soon he spotted a bearded seal pup sleeping on fl oating ice and signaled to his cousin, who killed it. Frank then saw its mother swimming nearby. His cousin told him not to hunt it but Frank didn?t want to give up. When the animal sur- faced directly in front of him, Frank took aim and fi red. I shot it immediately and hit it. I quickly harpooned it. It was very large. My poor heart was going tung-tung, and I could hear my heartbeat with my ears. I was afraid I would capsize, but I really wanted to catch it. After checking the ice, I got off on a thick spot, holding [the seal] with the harpoon. It was fl oating because it was very fat. I told my cousin Mancuaq to come, and he helped me. Qallaq didn?t make a sound again. We pulled it on top of the ice together and butchered it. Because it was so large, I told him to put it inside his kayak. I carried only the meat. When we got back, they came right over. Qilkilek didn?t say anything. . . . They butchered it, cutting it in strips, and distributed them. That?s what they do with fi rst catches. FIGURE 2. An uncatalogued hand-tinted lantern slide: ?Men hunt- ing in kayaks, probably near Kwigillingok,? in 1935/1936, Alaska, by Leuman M. Waugh, DDS, based on comments from Frank Andrew, 2002. Photo L02230 courtesy of National Museum of the American Indian, Smithsonian Institution. 07_Fienup-Riordan_pg079-088_Pole83 8307_Fienup-Riordan_pg079-088_Pole83 83 11/17/08 8:36:31 AM11/17/08 8:36:31 AM 84 SMITHSONIAN AT THE POLES / FIENUP-RIORDAN He asked me what I came for and told me to go back up and be the dumper of their urine buckets. They said they do that to those who they were encouraging because they want their minds to be stronger, those who they goaded toward success. Frank?s narrative is an unprecedented account, not only of what took place, but of the mixture of excitement, humil- ity, and determination he felt when ridiculed by men whose intent was not to shame him but to encourage him and prevent him from feeling overly proud and self- confi dent. Personal accounts are hard to elicit in gatherings and inter- views. Objects and photographs have opened doors that all of my questions never could. VISUAL REPATRIATION: ?EVERYTHING THAT IS MADE CAUSES US TO REMEMBER? To the extent that elders were personally moved by what they saw in collections, they regretted that young people in Alaska could not share their experience. Elders agreed that people cannot understand what they do not see. Frank Andrew (August 2003) spoke from personal experience: ?Among the things I saw in the museum, I didn?t know about things that I had not used. That?s how our young people are. They don?t know what they haven?t seen.? Neva Rivers (March 2004) of Hooper Bay agreed: Even though they hear about them with their ears, they can- not replicate these things if they don?t see them. But if our young people see what they did long ago, they will understand. They cannot come [to museums] because they are too far away, but if they bring things to Alaska, they can replicate some that they want to continue to be seen. Again and again elders said how valuable it would be for young people to see what they were seeing. During our 2003 trip to MSC, Frank Andrew (February 2003) remarked: Only education can keep things alive, only if a person who listens closely hears the information. These handmade items were constantly constructed when I was young by those who knew how to make them and who had the knowledge. Men taught young men how to catch animals. These [tradi- tional ways] were visible and did not change. Today [these ways] are no longer displayed, because we are not teaching what we were taught, even though we have the knowledge. It will not live on if it is like that. Paul John (January 2004) agreed, ?We are losing our way of life, and we need to help young people and others to better understand what they?ve lost. If the things that our ancestors used are shown, they will think, ?So this is what our ancestors did, and I can do what my ancestors did, too.?? Statistics bear out elders? view that contemporary young people lack knowledge of and appreciation for the values and technical skills that made life both possible and meaningful on the Bering Sea coast in the not-so-distant past. Southwest Alaska has one of the highest suicide rates in the nation, primarily young men and women in their twenties. Rates of poverty, alcohol abuse, and domestic vi- olence are also disproportionately high. The rapid changes before and after Alaska statehood in 1959 shook the moral foundation of Yup?ik community life to its core. These problems run deep, and knowledge and pride in their past is one among many elements needed for a solution. Joan Hamilton (November 2003) of Bethel said simply, ?Many people here are displaced by alcohol, but if we learn about ourselves from our elders, our minds will improve.? The truth of Frank Andrew?s and Paul John?s words was brought home to me personally in April 2003 when I listened to Jeffery Curtis, a Toksook Bay high-school student, speak publicly about his recent visit to Anchor- age. He said how glad he was to have the opportunity to visit the University of Alaska that he hoped someday to attend. He said that he planned to study science because his ancestors had no science, and he wanted to learn what white people could teach. Jeff comes from a proud and talented family, and his grandfather, Phillip Moses, is a master kayak builder with expert knowledge on many as- pects of Yup?ik technology. Jeff knows this, but nowhere has he learned to respect his grandfather?s knowledge as ?science.? Phillip Moses had likewise given me food for thought six months before, during work in collections at the An- chorage Museum. I had undergone retinal surgery two weeks earlier and could still only half-see out of one eye. As we worked together, my partner Alice Rearden handed Phillip a pair of wooden snow goggles, painted black on the inside with long, thin slits to let in the light. Phillip smiled, passed them to me to examine, and then launched into an enthusiastic explanation of how these goggles were the original ?Yup?ik prescription sunglasses.? Half- listening, I held the goggles to my eyes, and for the fi rst time since surgery, I could see! As I digested the sophisti- cated design? thin slits that focused the light like a pinhole camera, enhancing the user?s vision? I could hear Phillip 07_Fienup-Riordan_pg079-088_Pole84 8407_Fienup-Riordan_pg079-088_Pole84 84 11/17/08 8:36:33 AM11/17/08 8:36:33 AM YUP?IK ESKIMO CONTRIBUTIONS TO ARCTIC RESEARCH 85 relating in Yup?ik how the goggles worked both to reduce glare and to help a hunter see far. Phillip, like many elders, was well aware of the goggles? properties, yet I know of no reference in the literature on southwest Alaska regarding the capacity of snow goggles to improve distance vision. Like Phillip, many living elders can articulate the funda- mentals of Yup?ik technology. How powerful it would be to bring their clear descriptions home to a younger genera- tion, both Native and non-Native. Finally, in 2003, the Calista Elders Council began to search for ways to respond to the desire of their board of elders to bring museum objects home. Repatriation was not the issue, as ownership of objects was not the goal. Rather, ?visual repatriation? was what they sought? the opportunity to show and explain traditional technology to contemporary young people. Just as the Yup?ik community had looked to the An- chorage Museum in 1993 when beginning work on the Yup?ik mask exhibit, it again turned to the museum, which energetically embraced their project. Planning meetings formally began in August 2003 with a combination of National Science Foundation and Anchorage Museum As- sociation support. The fi rst meeting took place in Bethel in August 2003. There, a team of twelve Yup?ik elders and educators? including Frank Andrew and Paul John? gathered to plan a comprehensive exhibit of nineteenth- century Yup?ik technology. First, we discussed what kinds of objects the Yup?ik community would want to see. The answer was ?every- thing.? This was no surprise, given the elders? all-inclusive choices three years before at NMAI. What followed did surprise me, although in retrospect it should not have. I spoke briefl y about the mask exhibit that many of us had worked on together ten years earlier, saying that since that exhibit had focused on Yup?ik spirituality, we could take this opportunity to focus on Yup?ik science. I said that this exhibit could be what Agayuliyararput/Our Way of Making Prayer was not. I was reminded politely but fi rmly that Yup?ik tools and technology were also ?our way of making prayer.? Yup?ik team members did not view their traditional technology and spirituality as separable, and a valuable contribution of our exhibit would be to show how their ancestors lived properly, without this separa- tion. Elsie Mather explained, ?Long ago our beliefs and our way of life weren?t seen as separate. But nowadays, they look at those two as separate. In this exhibit, we should remember that and try to help people understand. If our exhibit becomes a reality, it will be taught that their ways of life and their beliefs were one.? Our second task was to name the exhibit. This was done with serious deliberation. After several suggestions, Frank Andrew spoke: ?The way of our ancestors is called yuungnaqsaraq [?to endeavor to live?]. When using all the tools together, only a person who is trying to survive will use them to live. That?s the name, and our ancestors used it all the time, ciuliamta yuungnaquciat [our ances- tors? way of life].? Paul John agreed: ?Back when Yup?ik people were really surviving on their own, they took care of themselves, trying to follow their traditions.? Mark John then added a crucial observation, restating the Yup?ik phrase in the present tense: We could make it more personal rather than distant. It could be yuungnaqpiallerput [the way we genuinely live], which in- cludes us, too. We are part of all that is being displayed. In the villages, people still utilize those ways, even though they may be using different materials. We?re not distancing ourselves from our ancestors. Paul John concluded: ?That yuungnaqpiallerput is perfect as a title. We really did try to live and survive the real way.? Discussion continued on which objects people thought most important to include. Paul John again mentioned the adze and the ax, as well as the fi re-making tools he had admired in New York. Frank Andrew spoke of the kayak and of that most essential tool, the negcik (gaff), which he referred to as ?life hook.? Andy Paukan remembered the powerful sinew-backed bow, and Marie Meade recalled the fi nely sewn clothing and ceremonial regalia she had seen in collections. Paul John emphasized the importance of including the drum as a metaphor for the continued vitality of the Yup?ik way of life. Frank Andrew concluded: ?The reverberation of the drum kept everyone together.? Elders also enthusiastically supported the inclusion of newly made examples of traditional technology, includ- ing a kayak, fi sh trap, seal-gut parka, and bearskin boat. Living elders had the skills to make these tools, and, once again, many people thought that elders mentoring young people in these techniques had the potential not only to transfer specifi c skills but also to shape lives. Another issue was how to organize the objects. A re- current theme was the continued importance of the sea- sonal cycle of activities, both in the past and today. They suggested that this cycle be used as the foundation for the exhibit. This simple but elegant mandate is what we have followed. Our story begins with preparation in the village and moves through spring, summer, fall, and early-winter 07_Fienup-Riordan_pg079-088_Pole85 8507_Fienup-Riordan_pg079-088_Pole85 85 11/17/08 8:36:33 AM11/17/08 8:36:33 AM 86 SMITHSONIAN AT THE POLES / FIENUP-RIORDAN harvesting activities. We then return to the winter village, where activities today, as in the past, focus on sharing the harvest and on renewal for the coming year. To tell this story, our exhibit includes examples of the most important features of nineteenth- and early- twentieth- century Yup?ik technology. It draws from a number of major collections of Yup?ik material culture in the United States and Europe, as well as from many less known but equally important collections. Some of our best pieces, however, come from the Smithsonian, including pieces col- lected by Edward Nelson, William Healey Dall, and A. H. Twitchell. Without Smithsonian collections, we could not tell our story. ?WE HAVE NO WORD FOR SCIENCE? In choosing a ?science? focus for their exhibition, Yup?ik community members continue to advocate for re- spect for their knowledge systems. The perceived gap be- tween Yup?ik indigenous knowledge and western science is enormous. Clearly, there are differences; but understand- ing the links can deepen our appreciation of both Yup?ik and western thought (Kawagley, 1995). When describing Yup?ik masks and ceremonies, elders made it clear that in the past they had no separate category for ?religion.? Everyday acts were equally ?our way of mak- ing prayer.? Similarly, discussions of hunting and harvesting activities make no separation between a person?s technical and moral education. Frank Andrew (February 2003) re- marked that ?Everything has a rule, no matter what it is. Because admonitions are a part of these snow goggles, we are talking about it through these.? Elsie Mather (Novem- ber 2003) observed, ?Our language had no word for sci- ence, yet our tools were so well designed that they allowed us to live in a land no one else would inhabit.? Yup?ik ontology promoted constant watchfulness and attention to the signs the natural world provided. A child?s fi rst task each morning was to exit the house and observe the weather. When traveling, each person depended for survival on observational skills honed from an early age. Knowledge in the past was situated, based on obser- vation and experience. Frank Andrew (June 2003) stated, ?I only speak intelligently about things that I know here in our village. I don?t know things in other villages that I didn?t see, and I cannot explain them very well.? What Frank does know, however, would impress any professional biologist or natural scientist. Frank and his contemporaries are gifted naturalists, engaged in classifi cation of all aspects of the world around them, often by appearance, usefulness, and behavior. Frank (June 2003) provided one excellent ex- ample. Previously he had talked at length about the differ- ent species of sea mammals, all of which have a one-to-one correspondence with western species classifi cations, includ- ing makliit (bearded seals), nayit (hair seals), issurit (spot- ted seals), qasrulget (ribbon seals), asveret (walrus), cetuat (beluga whales), arveret (bowhead whales), arrluut (killer whales), and arrnat (sea otters). He also distinguished be- tween different age groups within individual species. For example, the general category of bearded seal (Erignathus barbatus) includes maklak or tungunquq (adult bearded seal), maklassuk (subadult bearded seal), maklacuk (adult bearded seal with a small body but the fl ippers and intes- tines of an adult), qalriq (bearded seal in rut), amirkaq (young bearded seal), maklassugaq (two-year-old bearded seal), and maklaaq (bearded seal pup). Speaking to Alice Rearden and his son, Noah, both of whom he assumed understood the names for seal spe- cies and age groups, Frank added another level of detail, naming eight distinct varieties of bearded seals based on appearance and behavior, three of which I quote below: There are many bearded seals, and they all have different names. Some are rare, like those that have long beards that curl up when released. When they come out of the water close by, it seems as though they are biting on something large with their beards curled, looking like balls. They call those bearded seals ungagciaret [from ungak, ?whisker?]. . . . Then there are bearded seals that swim on their backs. When they get to the ice, they climb up face down, gallop across, go into the water, and then reappear on their backs. They say the ones that get sleepy do that. They said that if we saw one of those we should follow it carefully. They said that it would climb on top of the ice after awhile and stop and sleep. They say to hunt it when it does that. They called those papangluat. Then they say that some bearded seals would sleep and wake. When they look at their surroundings, they would curl up sitting on their stomachs with their head and hind fl ippers touch- ing, turning all the way around, looking behind them, searching their surroundings. After they look all around, they fi nally lie down and sleep. They call those ipuuyulit [from ipug-, ?to move with one?s front high in the air?]. They suddenly awake and search their surroundings. They are more afraid of the area behind them. That?s why they say not to approach them from behind, only by looking straight at them. We would approach them with them watching us. Close observation and classifi cation of the natural world are not the only things Yup?ik experts have in common with their western counterparts. At the same time elders reported 07_Fienup-Riordan_pg079-088_Pole86 8607_Fienup-Riordan_pg079-088_Pole86 86 11/17/08 8:36:34 AM11/17/08 8:36:34 AM YUP?IK ESKIMO CONTRIBUTIONS TO ARCTIC RESEARCH 87 important instructions that guided life, they tested rules as a means of judging their veracity. Rather than showing blind obedience to a timeless canon, Yup?ik men and women frequently describe their questioning of the principles on which they based their actions and understandings. Nick Andrew (March 2004) of Marshall described testing the admonishment that broad whitefi sh would become scarce if those caught in lakes were fed to dogs: I went with my male cousin when he was a boy to check our net with dogs. We got to the net and pulled it, and there were so many broad whitefi sh, and we set the net again. When we fi nished, I told him, ?Don?t tell on me, cousin.? I took them and gave one to each of the dogs. When they were done eating, we returned home. I told him, ?I wonder how our net will do tomor- row. Come with me again.? Then the next day, we checked our net. We pulled it, and it was heavy. We saw that we caught more than before. [Whitefi sh] don?t become scarce in lakes since they stay and don?t have anywhere to go. But they warned us not to throw them around or discard them carelessly. Yup?ik experimentation extended to technology. Men and women learned to construct and work with tools through constant trial and error. Kwigillingok elder Peter John (February 2003) noted: We tried to learn to make things. We took them by ourselves and examined them. We Yupiit are like that. We listen to and watch those who are working. Sometimes when we try to work, we don?t do a good job and stop working on it. When we try the next time, it looks better. Then we repeatedly make other ones. We just don?t do it once. That is the way to learn. Working in museum collections, one cannot fail to be impressed by the varied tool types and clothing patterns Yup?ik men and women created. There was a tool for ev- ery purpose. When Western technology was introduced, Yup?ik craftsmen embraced many labor-saving devices. If a new tool broke, time-tested materials were often used to fi x it, as when a commercially made boat propeller was re- placed by one fashioned from bone. The late Jim VanStone went so far as to dub Inuit peoples ?gadget ridden.? They knew their materials well and displayed impressive inven- tiveness in using them to advantage. Trial and error played a central role in Yup?ik learning and discovery. The perspectives shared by elders show important dif- ferences from and similarities with western science. Yup?ik knowledge was and is geared primarily to functions and outcomes. It is critical to know how to achieve some specifi c end so that resources necessary for survival and well-being may be acquired effectively. Western science is primarily aimed at developing and testing hypotheses to understand what is happening within and between vari- ables. However, the two are complementary in that Yup?ik science is the result of signifi cant trial and error that has produced acceptable outcomes, while western science can explain how these outcomes were achieved. Yup?ik technology can demonstrate scientifi c prin- ciples in new and exciting ways by matching such prac- tical outcomes to the phenomena they were designed to address. Moreover, the fact that Yup?ik science produced such outcomes prior to their conceptual bases is critical in understanding how ?science? as a process must be care- fully evaluated both on hypothesis testing and on mani- fested outcomes. Such a collaboration is especially impor- tant for science education in Alaska, where it can make the subject more relevant and effective. At our last exhibit-planning meeting, steering commit- tee members articulated the purpose of the exhibit in their own words. Elsie Mather stated, ?It will show the proven ways of tools and processes Yup?ik people used to survive and let people see the common ways we share the knowl- edge of our environment.? Joan Hamilton said, ?It will help people understand science and how it is part of everyday life, and it will communicate how much knowledge of the world Yup?ik people needed to survive.? There is a great deal of misunderstanding regarding how science works as a process unto itself. The value of considering western sci- entifi c approaches side by side with those of Yup?ik tradi- tional knowledge to close the gap between academic venues and the general public cannot be overstated. Yup?ik grade- school principal Agatha John (March 2004) of Toksook Bay articulated the dilemma of her generation: ?When I was in school I hated science. I couldn?t understand it. Not only was it in another language [English], but all the examples were foreign. If we begin to speak of ?Yup?ik science,? we will give our children something they can understand.? In closing, I would like to return to Yup?ik motiva- tions for traveling to the Smithsonian, sharing informa- tion, and seeking to borrow objects to display in Alaska and beyond during this Fourth International Polar Year. They hope to create an exhibition that will teach about the Yup?ik way of life? the animals and plants they rely on, the tools they used to survive, and the values that ani- mate their lives. Perhaps more important, their work at the Smithsonian teaches us about the generosity and com- passion of men and women who shared their knowledge, not only to inform us but to enrich all our lives and allow us to live genuinely. 07_Fienup-Riordan_pg079-088_Pole87 8707_Fienup-Riordan_pg079-088_Pole87 87 11/17/08 8:36:34 AM11/17/08 8:36:34 AM 88 SMITHSONIAN AT THE POLES / FIENUP-RIORDAN ACKNOWLEDGMENTS I am indebted fi rst and foremost to the Yup?ik men and women who generously shared their knowledge, and to Calista Elders Council (CEC) language expert Alice Rearden for carefully and eloquently transcribing and translating what they said. Our work together in Alaska has been supported by the CEC with funding from the National Science Foundation. We are grateful to the many Smithsonian staff members, both at the National Museum of Natural History and at the National Museum of the American Indian, who made it possible for Yup?ik elders to work in their collections. Special thanks to Bill Fitz- hugh, Smithsonian Institution, for fi rst bringing me into the Smithsonian in the 1980s, and to both Igor Krupnik, Smithsonian Institution, and Bill Fitzhugh for the invita- tion to share what I learned. NOTE ABOUT INTERVIEW DATES Dates following Elders? names refer to the month and year discussions with them took place. Tapes are archived with the Calista Elders Council, Bethel, Alaska. LITERATURE CITED Fienup-Riordan, Ann. 1996. The Living Tradition of Yup?ik Masks: Agayuliyararput/Our Way of Making Prayer. Seattle: University of Washington Press. ???. 2005. Yupiit Qanruyutait/Yup?ik Words of Wisdom. Lincoln: University of Nebraska Press. ???. 2007. Yuungnaqpiallerput/The Way We Genuinely Live: Master- works of Yup?ik Science and Survival. Seattle: University of Wash- ington Press. Fitzhugh, William W., and Susan A. Kaplan. 1982. Inua: Spirit World of the Bering Sea Eskimo. Washington, D.C.: Smithsonian Institution Press. Kawagley, Oscar. 1995. A Yupiaq Worldview: A Pathway to Ecology and Spirit. Prospect Heights, Ill.: Waveland Press. Krupnik, Igor, and Stephen Loring. 2002. The Waugh Collection Proj- ect: ASC Joins Efforts with the National Museum of the American Indian. ASC Newsletter, 10: 24? 25. Nelson, Edward William. 1899. The Eskimo about Bering Strait. Bureau of American Ethnology Annual Report for the Years 1896? 1897, Volume 18, Part 1. Washington, D.C.: Smithsonian Institution Press. 07_Fienup-Riordan_pg079-088_Pole88 8807_Fienup-Riordan_pg079-088_Pole88 88 11/17/08 8:36:34 AM11/17/08 8:36:34 AM ABSTRACT. From 1881 to 1883, as part of the First International Polar Year, an ex- pedition sponsored by the U.S. Signal Corps and the Smithsonian Institution operated a research station a short distance north of where the modern city of Barrow now stands. The 10 members of the expedition had the primary task of making an unbroken series of weather and magnetic observations over the two-year period, and the secondary task of studying the natural history of the Barrow area. ?Natural history? included descriptions of native life and collections of material culture, in addition to studies of the fauna and fl ora. In this paper, I summarize the substantial contributions to our knowledge of North Alaskan Eskimo life made by members of the expedition, and evaluate them in the light of work that has been done since. INTRODUCTION From 1881 to 1883, as part of the fi rst International Polar Year, an expedi- tion sponsored jointly by the U. S. Signal Corps and the Smithsonian Institu- tion operated a research station near Point Barrow, Alaska. 1 The members of the expedition had the primary task of making an unbroken series of weather and magnetic observations over the two-year period, and the secondary task of studying the natural history of the Barrow area. ?Natural history? was under- stood to include descriptions of the local people and collections of their mate- rial culture, in addition to observations of the fauna and fl ora. In this paper, I summarize and contextualize the contributions they made to our knowledge of North Alaskan I?upiaq Eskimo life. Point Barrow is the northernmost point of Alaska and of the United States as a whole. It is approximately 550 kilometers north of the Arctic Circle, and 400 kilometers north of the latitudinal tree line (Figure 1). It is located in the Beaufort coastal plain ecoregion, a treeless area of very low relief having a con- siderable amount of surface water (Nowacki et al., 2002). Summers are short and cool, and the winters are long and cold. The climate was signifi cantly colder in the nineteenth century than it is now. During the winter, the nearby ocean was completely frozen over; for much of the summer, it was covered with unconsoli- dated fl oating ice. Ernest S. Burch Jr., 3601 Gettysburg Road, Camp Hill, PA 17011-6816, USA (esburchjr@aol.com). Accepted 9 May 2008. Smithsonian Contributions to Alaskan Ethnography: The First IPY Expedition to Barrow, 1881? 1883 Ernest S. Burch Jr. 08_Burch_pg089-098_Poles.indd 8908_Burch_pg089-098_Poles.indd 89 11/17/08 8:31:57 AM11/17/08 8:31:57 AM 90 SMITHSONIAN AT THE POLES / BURCH EARLY EXPEDITIONS The fi rst westerners to visit the Point Barrow district were the members of a detachment from the Frederick W. Beechey expedition led by Thomas Elson, which arrived from the southwest in September 1826 (Beechey, 1831:414? 442). The explorers were met with a friendly greeting from the Natives, but that was quickly followed by considerable hostility. Not only for that reason, but also because the sea- son was dangerously far advanced, the men turned around and headed back south almost immediately. The second western expedition to make contact with the Native people of Point Barrow was a detachment from a Hudson?s Bay Company expedition sent to explore the western Arctic coast of North America. The group was led by Thomas Simpson, and it arrived at the point from the east on August 4, 1837 (Barr, 2002:70? 112; Simpson, 1839; 1843). Simpson?s small party was fortunate to get there when it did, because the settlement was largely un- occupied at the time. The few residents who were there were frightened and hid from the explorers. However, they were soon persuaded to come out and show themselves. Everyone got along pretty well for the few hours that the explorers were there. The next year, on July 23, 1838, a Russian expedi- tion led by Aleksandr Kashevarov reached Point Bar- row in a fl eet of small boats, arriving from the southwest ( VanStone, 1977:31? 45). A fair number of people, with FIGURE 1. Map of Alaska. 08_Burch_pg089-098_Poles.indd 9008_Burch_pg089-098_Poles.indd 90 11/17/08 8:31:57 AM11/17/08 8:31:57 AM SMITHSONIAN CONTRIBUTIONS TO ALASKAN ETHNOGRAPHY 91 considerable hostility, met Kashevarov?s party. The Rus- sians were forced to fl ee in fear of their lives after just three days. Despite the brevity of his stay, Kashevarov acquired some very useful information on native life in the Barrow district. This was due to the fact that his party included an interpreter. Kashevarov is the only person, Native or otherwise, who has ever reported the name of the nation (Burch, 2005:11? 33) whose members inhabited the Bar- row district. According to what he was told, they called themselves, and were known by others, as ?Kakligmiut? (VanStone, 1977:33). FRANKLIN SEARCH EXPEDITIONS The fourth western expedition to visit Point Barrow consisted of several of the ships involved in the search for the lost British explorer Sir John Franklin (Bockstoce, 1985). One of them, the depot ship H. M. S. Plover, under the command of Rochefort Maguire, spent the winters of 1852? 1853 and 1853? 1854 frozen in the ice a short dis- tance southeast of Nuvuk, the I?upiaq settlement on the point (Figure 2). The British were greeted with consider- able hostility. However, through wise diplomacy by the leaders on both sides, peaceful relations were established, and were maintained for the rest of the time the British were there. The information on native life acquired by the members of this expedition exceeded that of its three pre- decessors by several orders of magnitude. The surgeon on the Plover, John Simpson, already had acquired some profi ciency in the I?upiaq language when the ship spent successive winters on Kotzebue Sound (1849? 1850) and in Grantley Harbor (1850? 1851), on the western end of the Seward Peninsula. He was assigned the task of learning about native life in the Barrow district, in addition to his duties as surgeon. He performed his re- search through almost daily contact with Nuvuk?s inhabit- ants; this was mostly when they visited the ship, but also through his periodic visits to the village. Simpson (1855; 1875) wrote an outstanding report on what he learned about native life. It was one of the best ethnographic accounts of any indigenous North American people to appear in the nineteenth century. Much more FIGURE 2. Former I?upiaq settlements in the Barrow district, Alaska. 08_Burch_pg089-098_Poles.indd 9108_Burch_pg089-098_Poles.indd 91 11/17/08 8:32:32 AM11/17/08 8:32:32 AM 92 SMITHSONIAN AT THE POLES / BURCH recently, John Bockstoce (1988) published an edited and annotated version of Captain Maguire?s diary of the years spent near Point Barrow. He included in the volume a re- print of Simpson?s 1855/1875 report and several other use- ful documents. More recently still, Simpson?s (1852? 1854) diary, as well as several other manuscripts written by him, became accessible in the Duke University Archives. These documents, plus others produced by people involved in the Franklin search (e.g., Collinson, 1889, Hooper, 1853, Maguire, 1857, Pim, 1853, Pullen, 1979, Seemann, 1853), contain a remarkable amount of information on native life in the Barrow district in the mid-nineteenth century. We know not only what the members of the expedition found out, but also, through the diaries, how and from whom many of them acquired their information. THE IPY EXPEDITION 2 The IPY expedition to Barrow arrived 27 years after the Plover left. The members of this expedition established a base on shore about 15 kilometers southwest of the point. It was near Utqiag? vik, the other main settlement of the ?Kakligmiut.? Its leader, Patrick Henry Ray, did not want to establish a base at Nuvuk because the only dry ground there was already taken by the native village. He also did not want to locate the base right in Utqiag? vik be- cause he was afraid of being pestered by the Natives. So, it was set up a little more than 1 kilometer to the northeast, at the place known more recently as Browerville. Some tension between the Natives and the research- ers arose due to the fact that the commander tried to put a stop to the trade in whiskey and fi rearms that was be- ing conducted with American whalers during the period when the expedition was based there (Ray, 1882b, 1882c). However, in general, the two groups got along pretty well. The commander, Patrick Henry Ray (1882a), wrote that ?these people in their appearance, general intelligence and industry are superior to any native I have seen on the conti- nent. . . .? His colleague, John Murdoch (1890a:223), said that the Eskimos were ?altogether pleasant people to see and to associate with.? Another colleague, Middleton Smith (1902:118), characterized them as ?a good people.? One does not expect to read such positive sentiments expressed by late-nineteenth century-American white men about in- digenous North Americans. They help account for the ex- pedition members? willingness to loan tools and sometimes weapons to their I?upiaq neighbors, at least during the sec- ond year of their stay. (I have seen no evidence on what the residents of Utqiag? vik thought of the IPY people.) The expedition members consisted of 10 men of whom fi ve are of special importance to this paper. In what follows I summarize the individual contributions made by these fi ve, plus one other person, and then discuss the expedition?s collective results. PATRICK HENRY RAY Patrick Henry Ray was a fi rst lieutenant in the 8th infantry. It is not quite clear to me just how he spent his time in Utqiag? vik. He was apparently not involved in the boring, time-consuming work of recording meteorologi- cal and magnetic observations. Instead, he managed to get out and about a fair amount of the time, both in the na- tive village and beyond. For example, in late March and early April of both 1882 and 1883, he traveled south to the Meade River with Native companions during caribou hunting season (Ray, 1988a:lii; 1988b:lxxvii). He also vis- ited the settlement on numerous occasions. There he was able to observe ceremonies and rituals, as well as people simply going about their daily lives (Murdoch, 1988:80, 432; Ray, 1988c:xciii). Ray wrote informative summaries of the expedition and of his own travels inland (Ray, 1988a, 1988b), as well as a comprehensive sketch of native life (Ray, 1988c). The latter is a generally accurate document, but it is dimin- ished by the fact that its author paraphrased and even pla- giarized the work of John Simpson from 30 years earlier. There is evidence (e.g., in Murdoch, 1988:433) that Ray kept a notebook, but if he did, it has been lost. E. P. HERENDEEN The second person worthy of mention is Captain Edward Perry Herendeen, a whaler and trader with con- siderable experience in northern Alaska ( J. Bockstoce, pers. comm.). Herendeen was brought along as inter- preter and storekeeper. Just how effectively he acted as an interpreter is questionable. Other members of the ex- pedition complained about being unable to communicate effectively with the Natives during the fi rst year of their stay (e.g., Murdoch, 1988:45), and Ray (1988c:lxxxvii) stated fl atly that the party had no interpreter. By the sec- ond year, each man could do fairly well on his own (Ray, 1988a:li). Herendeen seems to have gotten out and about even more than Ray did. He attended a number of ceremonies in the village, hunted inland with Natives in both fall and winter, and visited the whaling camps on the sea ice in spring. Others (e.g., Murdoch, 1988:39, 272, 276, 364, 08_Burch_pg089-098_Poles.indd 9208_Burch_pg089-098_Poles.indd 92 11/17/08 8:33:01 AM11/17/08 8:33:01 AM SMITHSONIAN CONTRIBUTIONS TO ALASKAN ETHNOGRAPHY 93 372, 374, 423) cite Herendeen as having provided them with information about a variety of subjects that he had to have obtained in the village or out in the country. If he kept a journal, it has been lost, and the only publication he produced that I am aware of was a piece on caribou hunting (Herendeen, 1892). 3 GEORGE SCOTT OLDMIXON George Scott Oldmixon was the surgeon on the ex- pedition. In addition to treating the health problems of expedition members, he also treated many sick Natives, and he made many visits to the village for that purpose. In the process, he learned something about native health problems and their means of dealing with them. Unfortu- nately, he, too, is not known to have kept a journal. How- ever, he reported some of what he learned to others, who wrote down some of what he told them. Oldmixon?s one substantive contribution to the expedition?s published re- ports was a set of height and weight measurements made of a number of I?upiaq men and women from the two Barrow villages (Murdoch, 1988:cvii). MIDDLETON SMITH Middleton Smith was one of the assistants who checked the instruments and kept the records of the mag- netic and meteorological observations. As far as I am aware, the only thing he wrote about the expedition was a popularized piece titled ?Superstitions of the Eskimo? (Smith, 1902). Unfortunately, the article contains some er- roneous information, such as the population fi gures on pp. 113? 114. It also paints an idyllic picture of I?upiaq life (e.g., on pp. 118? 119), contributing to the stereotype of Eskimos as being happy, hard-working, fun-loving, peace- ful people. In fact, like most people everywhere, they were considerably more complicated than that. Smith also presents interesting bits of information not included in the reports of his colleagues. For example, on page 120 he reports: When a death occurs in the village the women are not al- lowed, from sunset to sunrise, either to make or repair garments or to do sewing [of] any kind, except in the most urgent cases, when the work must be done while sitting within circles inscribed by the point of a knife upon the fl oor of the iglu. This is the only source I am aware of that reports on a way of circumventing a taboo. It would be fascinating to know if there were others. Also of interest are Smith?s ac- counts (pp. 127? 128) of how the Natives cheated during some of their trading sessions. JOSEPH S. POWELL Joseph S. Powell was not a member of the IPY expedi- tion, but he commanded the ship sent to re-supply it for the second winter. Powell?s (1988) report summarizes his visit, and also contains quite a bit of interesting informa- tion about native life in Utqiag? vik. Very little of this in- formation could have been obtained fi rst-hand, however. Powell was at Utqiag? vik for only a week, and at least some of the time was prevented from going ashore due to bad weather. Presumably, he also spent some of his time in su- pervisory activities onboard the ship. Rather than basing his account on personal observa- tion and experience, Powell relied on information con- veyed to him by others, primarily Lieutenant Ray and Ser- geant James Cassidy (Powell, 1988:lx). This had to have been presented to him in summary form rather than in de- tail, and, as a result, his report contains some useful gen- eralizations. For example: ?There are leading men whose infl uence depends on their wealth and the number of their relatives and friends, but no chiefs, hereditary or other- wise. . .? (Powell, 1988:lxi). I have never seen the subject of I?upiaq leadership characterized more accurately or succinctly than that. JOHN E. MURDOCH This brings us to John E. Murdoch. A Harvard-trained naturalist, Murdoch was one of the men involved in the te- dious recording of magnetic and meteorological data. Ac- cordingly, he was not inclined to go out as much as some of his colleagues. However, it is clear from comments scat- tered about in his various writings that he was interested in and generally aware of what was going on in Utqiag? vik and the area around it. He became profi cient enough in the I?upiaq language for people living on Norton Sound in western Alaska, to identify him later, on the basis of his speech, as someone coming from Point Barrow (Murdoch, 1988:46). Unlike John Simpson in the 1850s, Murdoch never reached a level of linguistic profi ciency at which he could discuss abstract philosophical matters, but he un- derstood enough to know that the Eskimos had a raun- chy sense of humor (Murdoch, 1988:419), and he could converse with them about a variety of day-to-day matters (e.g., Murdoch, 1988:58, 79, 384, 412, 424, 432). In addition to his other duties, Murdoch was put in charge of cataloguing the numerous artifacts that were 08_Burch_pg089-098_Poles.indd 9308_Burch_pg089-098_Poles.indd 93 11/17/08 8:33:02 AM11/17/08 8:33:02 AM 94 SMITHSONIAN AT THE POLES / BURCH obtained from the Natives through barter. He wrote the catalogue of ethnological specimens and the natural history sections of the expedition?s fi nal report (1885a, 1885b), and he authored a major monograph (1988) and more than a dozen articles. The latter concerned such varied subjects as fi sh and fi shing (1884), seal hunting (1885d), sinew-backed bows (1885e), legends (1886), native clothing and physique (1890a), counting and measuring (1890b), I?upiaq knowl- edge of heavenly bodies (1890c), whale hunting (1891), and I?upiaq knowledge of local wildlife (1898). DISCUSSION The primary objectives of IPY-1 were in the fi elds of physics and meteorology. Thus, as noted by Igor Krupnik (2009, this volume), 4 it is a curious fact that the most enduring products of the expedition to northern Alaska were its ethnographic collections and reports. Just why the members of this expedition engaged in studies of human affairs at all is not immediately clear. Of the main con- tributors in this area, Patrick Henry Ray, was a military man and John Murdoch was a naturalist, although both seem to have been intelligent and intellectually curious individuals. The answer to this question must lie in the considerable involvement of Spencer F. Baird, head of the Smithsonian Institution, in the expedition?s planning and staffi ng. Baird was a biologist with wide-ranging interests, he was an avid collector personally, and, at the time, he was engaged in an ambitious program to build up the mu- seum?s collections (Henson, nd). In 1879 he oversaw the integration of the Bureau of American Ethnology with the Smithsonian, and, ?with strong support from Congress, [he] encouraged the ethnologists to also collect artifacts and pursue archaeological investigations? (Henson, nd). Thus, it seems reasonable to conclude that Baird strongly encouraged/required the members of the expedition, par- ticularly Murdoch, to conduct ethnological/ethnographic research and to bring what they acquired back to the mu- seum. If the expedition was sent out with a set of specifi c ethnographic research objectives, however, no record of what it was has been discovered. We are thus forced to surmise that the various sections of Murdoch?s (1988) monograph refl ect a set of more or less ad hoc conceptual categories that enabled him to organize into a coherent account his own experiences and observations, as well as those reported to him by his colleagues. Nearly all pieces of the artifact collection were ob- tained through barter, with ?the natives bringing their weapons, clothing and other objects to the station for sale? (Murdoch, 1988:19). Since Murdoch was the person charged with recording these items, this gave him ?espe- cially favorable opportunities for becoming acquainted with the ethnography of the region? (Murdoch, 1988:19), even though he was not able to leave the station nearly as often as some of his colleagues. Murdoch apparently hoped to do more than just collect artifacts. The following passage expresses his frustration: It was exceedingly diffi cult to get any idea of the religious belief of the people, partly from our inability to make ourselves understood in regard to abstract ideas and partly from ignorance on our part of the proper method of conducting such inquiries. For instance, in trying to get at their ideas of a future life, we could only ask ?where does a man go when he dies?? to which we, of course, received the obvious answer, ?to the cemetery!? (Murdoch, 1988:430) Another passage elaborates: Occupied as our party was with the manifold routine scien- tifi c work of the station, it was exceedingly diffi cult to get hold of any of the traditions of the Natives, though they showed no unwillingness, from superstitious or other reasons, to talk freely about them. In the fi rst place there were so many (to the Eskimos) more interesting things to talk about with us that it was diffi cult to bring the conversation round to the subject in question. Then our lack of familiarity with the language was a great hindrance to obtaining a connected and accurate version of any story. The jargon, or kind of lingua franca, made up of Eskimo roots and ?pi- geon English? grammar, which served well enough for every-day intercourse with the Natives, enabled us, with the help of expres- sive gestures, to get the general sense of the story, but rendered it impossible to write down an Eskimo text of the tale which could afterwards be translated. Moreover, the confusion and diffi culty was still further increased by the fact that two or three people gen- erally undertook to tell the story at once. (Murdoch, 1886:594) The above factors resulted in the fact that the main ethnographic contribution of the IPY expedition lay in its collection of material objects, not in accounts of Native social organization, history, philosophy, or worldview. In the latter areas, John Simpson?s (1855; 1875) report remains the best single source, although there are many bits and pieces of new and updated, information in the IPY documents. However, the IPY collection of artifacts was signifi cant, the largest ever acquired in Arctic Alaska. Murdoch?s massive volume, fi rst published in 1892, is a superb adjunct to the collection itself because it provides excellent illustrations and descriptions of the objects and 08_Burch_pg089-098_Poles.indd 9408_Burch_pg089-098_Poles.indd 94 11/17/08 8:33:02 AM11/17/08 8:33:02 AM SMITHSONIAN CONTRIBUTIONS TO ALASKAN ETHNOGRAPHY 95 tells how many of them were made. It also compares items in the Barrow collection with similar artifacts acquired in other parts of the north by other expeditions. One important contribution the IPY reports in general made was to provide evidence of changes that had occurred in the Barrow people?s way of life during the 30 years since the Franklin Search Expedition. Perhaps the most striking difference was demographic. In 1853, the combined popu- lation of Nuvuk and Utqiag? vik had been about 540; 30 years later, it was less than 300 (Ray, 1988c:xcix). The intervening years had seen the arrival of American whaling ships and trading vessels in Arctic Alaskan waters. Many of these ships stopped briefl y at one or both of the Barrow villages almost every year after the Franklin Search Expedition left. The Americans brought fi rearms, ammuni- tion, whisky, and epidemic diseases to the Natives. They also killed a substantial number of the bowhead whales, on which the coastal native economy was based. Other changes resulted from the use of fi rearms in hunt- ing. Previously, seal hunters had fi rst attached themselves to a seal with a harpoon and a line, and only then killed the animal. With fi rearms, they killed the seal at a distance, then tried to attach a line to it for retrieval (Murdoch, 1885c). Whereas before they rarely lost a seal that had been struck, they now lost a signifi cant number, particularly in spring. Caribou hunting was also transformed. In the 1850s, Barrow hunters killed caribou in winter by digging pitfalls in the snow and killing the animals that fell into them. Since the snow was not deep enough in the fall to per- mit this, they did not hunt caribou at that time of year. By the 1880s, they could kill caribou at a distance with fi rearms, so they could hunt them in both fall and winter. This nearly doubled the hunting pressure on this particu- lar resource. POST-IPY RESEARCH The IPY expedition took place at a time when native life in the Barrow district was beginning to come into in- creasing contact with members of the U. S. Revenue Ma- rine and with an assortment of adventurers, explorers, and traders. Some of the individuals involved wrote informa- tive descriptions of native life in the region. However, none of them conducted systematic research and none of them made any effort to relate their observations to those of the earlier IPY or Franklin Search reports (of which they prob- ably were unaware.) This situation did not change until the 1950s, when some more serious investigations were undertaken. The major people involved in this subsequent work were Robert F. Spencer, Joseph Sonnenfeld, and Bar- bara Bodenhorn. ROBERT F. SPENCER The fi rst researcher to build on the work of the nineteenth-century investigators was Robert F. Spencer. 5 Spencer did his research in Barrow in the early 1950s, pub- lishing his fi ndings in the late 1950s, and for many years afterwards (e.g., 1959, 1967? 1968, 1968, 1972, 1984). Despite the late date of his fi eld studies, the emphasis in his writing was on the ?traditional? way of life, with the timeframe being left unspecifi ed. Careful examination of both his publications and his fi eld notes indicates that the situation he wrote about was what his informants experi- enced as children, in the late 1880s and 1890s ( Bodenhorn, 1989:24 n. 19). It certainly was not what J. Simpson and Maguire described for the early 1850s. Thus, even though Spencer did his research some seven decades after the IPY expedition left the fi eld, it is almost as though he did it just a few years later. Spencer fi lled in two major gaps left by his predeces- sors. First, he paid almost as much attention to I?upiat living inland as he did to those living along the coast (1959:3? 4, 132? 139). It seems hard to believe in retro- spect, but until Spencer?s book appeared, most anthro- pologists believed that Eskimos were primarily or even exclusively a coastal people. In fact, as Spencer (1959:21) pointed out, in the nineteenth century, inlanders outnum- bered coast dwellers in Arctic Alaska by a ratio of about three to one. He then went on to describe (1959:62? 97) in some detail the nature of the relations between the resi- dents of the two ecological zones, showing how they were linked into a larger regional system. Spencer?s second main contribution was to give the I?upiaq family system the attention it deserves (1959:62? 96; 1967? 68; 1968). Again, it seems hard to believe in retrospect, given the importance of families in I?upiaq so- cieties, but systematic studies of Eskimo family life were all but nonexistent at the time Spencer did his research. Spencer changed all that, and the decades following pub- lication of his major monograph witnessed an outpour- ing of kinship studies in Eskimo settlements all across the North American Arctic (Burch, 1979:72). JOSEPH SONNENFELD Joseph Sonnenfeld is a geographer who did research in Barrow for four months in 1954, the year after Spen- cer left. He apparently did not know of Spencer?s work 08_Burch_pg089-098_Poles.indd 9508_Burch_pg089-098_Poles.indd 95 11/17/08 8:33:03 AM11/17/08 8:33:03 AM 96 SMITHSONIAN AT THE POLES / BURCH at that time, and when he completed his Ph.D. thesis in 1957, he was acquainted only with Spencer?s report to the granting agency. As a result, he recapitulated some of Spencer?s reconstructive work. However, he was oriented much more to contemporary events than Spencer was. Thus, without really being aware of the fact, he brought the documentation of Barrow I?upiaq life forward from the end of the nineteenth century to the middle of the twentieth. Being a geographer, Sonnenfeld was interested primarily in ecological and economic matters. With reference to those subject areas, his Ph.D. thesis (1957) was wide ranging and informative. Unfortunately, it was never published. As far as I am aware, Sonnenfeld published only two articles on his work in Barrow, one (1959) on the history of domesti- cated reindeer herds in the Barrow district, the other (1960) on changes in Eskimo hunting technology. BARBARA BODENHORN Barbara Bodenhorn is a social anthropologist who be- gan her research in Barrow in 1980 and who continued it for many years subsequently. More than any of her pre- decessors, Bodenhorn tried to learn how the community worked as a social system, and she spent enough time in it to fi nd out. Her work focused on families, the inter- relations between and among families, and the role of fami- lies in the overall economy. She has written extensively on these and related subjects (e.g., 1989, 1990, 1993, 1997, 2000, 2001). Bodenhorn?s work is only the most recent effort in more than a century and a half of ethnographic research in Barrow. No other Arctic community has been so thor- oughly studied over so long a time. Viewed from this broad perspective, although their artifact collection remains un- equaled, the ethnographic work of John Murdoch, Patrick Henry Ray, and the other IPY expedition members con- stitutes just one link in a long chain of empirical investi- gations. The ?chain? as a whole should now become the center of someone?s attention: Where else in the Arctic can one fi nd so much good information on social change in one community over so long a time? CONCLUSION In conclusion, I wish to address briefl y the issue of what scientifi c value there might be in gaining knowledge of nineteenth-century native life in Barrow. The answer lies in the value of natural experiments. Social scientists can experiment with small numbers of people in highly restricted settings, but there is no way that we can experiment with entire societies, certainly not on any kind of ethical basis. The only way we can develop a broad understanding of how human social systems oper- ate is by observing people going about their lives in their own ways without any interference from a researcher. This is what is ?natural? about the method. The greater the diversity of the social systems that can be studied in this way, the more ?experimental? the approach becomes, and the more powerful any resulting theories about the struc- ture of human social systems are likely to be. Arctic peoples in general, and Eskimos in particular, are important in this regard because they lived in such ex- treme environments. In the nineteenth century, the way of life of the Eskimos in the Barrow district stood in marked contrast to the previously recorded ways Eskimos lived in the eastern North American Arctic. This information ex- pands our knowledge of the range of variation of Arctic social systems and, by extension, of human social systems in general. In order to be scientifi cally signifi cant, though, the research on the societies in a sample of societies must be conducted in terms of a common conceptual and theo- retical framework so that a systematic comparative analy- sis can be subsequently carried out. The Franklin Search and IPY-1 reports are fairly close to meeting that require- ment, but they are only a fi rst step. Fortunately, the infor- mation they contain is good enough and complete enough for future researchers to adapt for that purpose. 6 NOTES 1. I thank John Bockstoce and Igor Krupnik for information and/or advice given during the preparation of this article. 2. Most of the expedition?s reports, originally published in the nineteenth century, were reprinted in 1988 by the Smithsonian Institu- tion Press. While both the earlier and later versions are listed here in the References, only the 1988 versions are cited in the text. 3. I have not seen this article myself, but I thought its existence should be recorded here. 4. Krupnik, this volume. 5. My knowledge of Spencer?s work was enhanced under a grant from the National Science Foundation, Offi ce of Polar Programs (OPP- 90817922). I am grateful to that organization for its support, and to Marietta Spencer for giving me her late husband?s Barrow fi eld notes. 6. A comparative analysis of the meteorological data acquired in IPY-1 was not carried out until recently (Wood and Overland, 2006). Thus, one of the primary objectives of the fi rst IPY was not achieved until nearly a century and a quarter after the raw data were collected. 08_Burch_pg089-098_Poles.indd 9608_Burch_pg089-098_Poles.indd 96 11/17/08 8:33:03 AM11/17/08 8:33:03 AM SMITHSONIAN CONTRIBUTIONS TO ALASKAN ETHNOGRAPHY 97 LITERATURE CITED Barr, William, ed. 2002. From Barrow to Boothia. The Arctic Journal of Chief Factor Peter Warren Dease, 1836? 1839. Montreal: McGill- Queen?s University Press. Beechey, Frederick W. 1831. Narrative of a Voyage to the Pacifi c and Bering?s Strait to Cooperate with the Polar Expeditions Performed in His Majesty?s Ship Blossom . . . in the Years 1825, 26, 27, 28. London: Colburn and Bentley. Bockstoce, John R. 1985. ?The Search for Sir John Franklin in Alaska.? In The Franklin Era in Canadian Arctic History, 1845? 1859, ed. Patricia D. Sutherland, pp. 93? 113. Mercury Series, Archaeologi- cal Survey of Canada Paper No. 131. Ottawa: National Museum of Man. ???, ed. 1988. The Journal of Rochfort Maguire, 1852? 1854: Two Years at Point Barrow, Alaska, aboard HMS Plover in the Search for Sir John Franklin. 2 vols. (2nd series, nos. 169 and 170). Lon- don: Hakluyt Society. Bodenhorn, Barbara A. 1989. ?The Animals Come to Me, They Know I Share?: I?upiaq Kinship, Changing Economic Relations and Endur- ing World Views on Alaska?s North Slope. 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Senri Ethnological Studies No. 53. Osaka, Japan: National Museum of Ethnology ???. 2001. ??He Used to Be My Relative?: Exploring the Bases of Relatedness among I?upiaq of Northern Alaska.? In Cultures of Relatedness: New Approaches to the Study of Kinship, ed. Janet Carstens, pp. 128? 148. Cambridge, U.K.: Cambridge University Press. Burch, Ernest S., Jr. 1979. The Ethnography of Northern North America. A Guide to Recent Research. Arctic Anthropology, 16(1): 62? 146. ???. 2005. Alliance and Confl ict: The World System of the I?upiaq Eskimos. Lincoln: University of Nebraska Press. Collinson, Richard. 1889. Journal of HMS Enterprise on the expedition in Search of Sir John Franklin?s Ships by Behring Strait, 1850? 1855. London: Sampson, Low, Marston, Searle and Rivington. Henson, Pamela M. n.d. Spencer F. Baird?s Vision for a National Mu- seum. http://www .siarchives .si .edu/ history/ exhibits/ baird/ bairdhm. htm (accessed 18 August 2008). Herendeen, E. P. 1892. An Esquimaux Caribou Hunt. Forest and Stream, 38(Mar. 17): 249. Hooper, William H. 1853. Ten Months among the Tents of the Tuski. London: John Murray. Krupnik, Igor. 2009. ??The Way We See It Coming?: Building the Legacy of Indigenous Observations in IPY 2007? 2008.? In Smithsonian at the Poles: Contributions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 129? 142. Washington, D.C.: Smithsonian Institution Scholarly Press. Maguire, Rochefort. 1857. ?Narrative of Commander Maguire, Wintering at Point Barrow.? In The Discovery of the North-West Passage. . ., 2nd edition, ed. Sherard Osborn, pp. 409? 463. Lon- don: Longman, Brown, Green, Longmans & Roberts. Murdoch, John. 1884. ?Fish and Fishing at Point Barrow, Arctic Alaska.? In Transactions of the American Fish Cultural Association, Thir- teenth Annual Meeting, pp. 111? 115. ???. 1885a. ?Catalogue of Ethnological Specimens Collected by the Point Barrow Expedition.? In Report of the International Polar Ex- pedition to Point Barrow, Alaska, ed. P. H. Ray, pp. 61? 87. Wash- ington, D.C.: Government Printing Offi ce. ???. 1885b. ?Natural History.? In Report of the International Polar Expedition to Point Barrow, Alaska, ed. P. H. Ray, pp. 89? 200. Washington, D.C.: Government Printing Offi ce. ???. 1885c. The Retrieving Harpoon: An Undescribed Type of Es- kimo Weapon. American Naturalist, 19: 423? 425. ???. 1885d. Seal-Catching at Point Barrow. Transactions of the An- thropological Society of Washington, 3: 102? 108. ???. 1885e. The Sinew Backed Bow of the Eskimo. Transactions of the Anthropological Society of Washington, 3: 168? 180. ???. 1886. A Few Legendary Fragments from the Point Barrow Eski- mos. American Naturalist, 20: 593? 599. ???. 1890a. Dress and Physique of the Point Barrow Eskimo. Popular Science Monthly, 38: 222? 229. ???. 1890b. Notes on Counting and Measuring among the Eskimo of Point Barrow, Alaska. American Anthropologist, 3: 37? 43. ???. 1890c. Notes on Names of the Heavenly Bodies and the Points of the Compass among the Point Barrow Eskimo. American An- thropologist, 3: 136. ???. 1891. Whale-Catching at Point Barrow. Popular Science Monthly, 38: 830? 836. ???. 1892. Ethnological Results of the Point Barrow Expedition. Ninth Annual Report of the Bureau of Ethnology for the Years 1887? 1888. Washington, D.C.: Government Printing Offi ce. (Re- printed 1988, Smithsonian Institution Press.) ???. 1898. The Animals Known to the Eskimos of Northwestern Alaska. American Naturalist, 32 (382):720? 734. ???. 1988. Ethnological Results of the Point Barrow Expedition. Smithsonian Classics of Anthropology Series reprint, No. 6. Wash- ington, D.C: Smithsonian Institution Press. Nowacki, Gregory, Page Spencer, Michael Fleming, Terry Brock, and Torre Jorgenson. 2002. Unifi ed Ecoregions of Alaska: 2001. U.S. Geological Survey Open-File Report 02? 297. Pim, Bedford. 1853. ?Journal.? In Narrative of the Voyage of H.M.S. Herald during the Years 1845? 51 in Search of Sir John Franklin, vol. II, by Berthold Carl Seemann, pp. 130? 148. London: Reeve and Company. Powell, Joseph S. 1988. ?Report of the Relief Expedition of 1882.? In Ethnological Results of the Point Barrow Expedition, pp. lv? lxiii. Smithsonian Classics of Anthropology Series reprint, No. 6. Wash- ington, D.C.: Smithsonian Institution Press. Pullen, H. F., ed. 1979. The Pullen Expedition in Search of Sir John Franklin: The Original Diaries, Log, and Letters of Commander W. J. S. Pullen. Toronto: Arctic History Press. Ray, Patrick Henry. 1882a. Letter, Ray to the Secretary, Offi ce of the Interior, from Barrow, May 18, 1882. Records of the International Polar Expedition to Point Barrow, Alaska. RG 27.4.6, box 5: cop- ies of letters sent, pp. 5? 6. U.S. National Archives II, College Park, Maryland. ???. 1882b. Letter, Ray to the Chief Signal Offi ce, from Barrow, July 23, 1882. Records of the International Polar Expedition to Point 08_Burch_pg089-098_Poles.indd 9708_Burch_pg089-098_Poles.indd 97 11/17/08 8:33:04 AM11/17/08 8:33:04 AM 98 SMITHSONIAN AT THE POLES / BURCH Barrow, Alaska. RG 27.4.6, box 5: copies of letters sent, pp. 11? 12. U.S. National Archives II, College Park, Maryland. ???. 1882c. Letter, Ray to the Chief Signal Offi ce, from Barrow, Au- gust 13, 1882. Records of the International Polar Expedition to Point Barrow, Alaska. RG 27.4.6, box 5: copies of letters sent. U.S. National Archives II, College Park, Maryland. ???. 1988a. ?Work at Point Barrow, Alaska, from September 16, 1881, to August 25, 1882.? In Ethnological Results of the Point Barrow Expedition, pp. l? liv. Smithsonian Classics of Anthropol- ogy Series reprint, No. 6. Washington, D.C.: Smithsonian Institu- tion Press. ???. 1988b. ?Narrative.? In Ethnological results of the Point Barrow expedition, pp. lxix? lxxxvi. Smithsonian Classics of Anthropology Series reprint, No. 6. Washington, D.C.: Smithsonian Institution Press. ???. 1988c. ?Ethnographic Sketch of the Natives of Point Barrow.? In Ethnological results of the Point Barrow expedition, pp. lxxx- vii? cv. Smithsonian Classics of Anthropology Series reprint, No. 6. Washington, D.C.: Smithsonian Institution Press. Seemann, Berthold Carl. 1853. Narrative of the Voyage of H.M.S. Herald during the Years 1845? 51 in Search of Sir John Franklin. 2 vols. Lon- don: Reeve and Company. Simpson, John. 1852? 1854. Point Barrow Journal, 1852? 1854. John Simpson Papers, Rare Book, Manuscript, and Special Collections Library, Box 5: Accounts of voyages. Oversized. Duke University, Durham, North Carolina. ???. 1855. Observations on the Western Esquimaux and the Country They Inhabit; From Notes Taken During Two Years at Point Barrow, by Mr. John Simpson, Surgeon, R.N, Her Majesty?s Discovery Ship ?Plover.? Great Britain. Parliament. House of Commons. Sessional Papers, Accounts and Papers 1854? 55 35, no. 1898: 917? 942. ???. 1875. Observations on the Western Eskimo and the Country They Inhabit. In A Selection of Papers on Arctic Geography and Ethnology, Reprinted and Presented to the Arctic Expedition of 1875, pp. 233? 275. (Reprint of 1855 report.) London: Royal Geo- graphical Society. Simpson, Thomas. 1839. Arctic Discovery. Nautical Magazine, 8: 564? 565. ???. 1843. Narrative of the Discoveries on the North Coast of Amer- ica: Effected by the Offi cers of the Hudson?s Bay Company during the Years 1836? 39. London: R. Bentley. Smith, Middleton. 1902. ?Superstitions of the Eskimo.? In The White World, ed. Rudolph Kersting, pp. 113? 130. New York: Lewis, Scribner, and Co. Sonnenfeld, Joseph. 1957. Changes in Subsistence among the Barrow Es- kimo. Ph.D. diss., Johns Hopkins University, Baltimore, Maryland. ???. 1959. An Arctic Reindeer Industry: Growth and Decline. The Geographical Review, 59(1): 76? 94. ???. 1960. Changes in an Eskimo Hunting Technology: An Introduc- tion to Implement Geography. Annals of the Association of Ameri- can Geographers, 50(2): 172? 186. Spencer, R. F. 1959. The North Alaskan Eskimo: A Study in Ecology and Society. Bureau of American Ethnology Bulletin 171. Washington, D.C.: Smithsonian Institution Press. ???. 1967? 1968. Die organization der Ehe under den Eskimos Nor- dalaskas. Wiener Vu?lkerkundliche Mitteilungen, ns 9/10: 13? 31. ???. 1968. ?Spouse Exchange among the North Alaskan Eskimo.? In Marriage, Family, and Residence, ed. Paul Bohannan and J. Middle- ton, pp. 131? 146. New York: Natural History Press. ???. 1972. ?The Social Composition of the North Alaskan Whaling Crew.? In Alliance in Eskimo Society: Proceedings of the American Ethnological Society, 1971, Supplement, ed. Lee Guemple, pp. 110? 131. Seattle: University of Washington Press. ???. 1984. ?North Alaskan Coast Eskimo.? In Handbook of North American Indians, ed. William C. Sturtevant. Volume 5: Arctic, ed. David Damas, pp. 320? 327. Washington, D.C.: Smithsonian Insti- tution Press. VanStone, James W., ed. 1977. A. F. Kashevarov?s Coastal Explorations in Northwest Alaska, 1838. Trans. David H. Kraus. Fieldiana: An- thropology, 69. Wood, Kevin R., and James E. Overland. 2006. Climate Lessons from the First International Polar Year. Bulletin of the American Meteo- rological Society, 87(12): 1685? 1697. 08_Burch_pg089-098_Poles.indd 9808_Burch_pg089-098_Poles.indd 98 11/17/08 8:33:04 AM11/17/08 8:33:04 AM ABSTRACT. In the northern Bering Sea and Arctic Ocean, a 2,000- year tradition of Alaska Native bowhead whaling continues to the present day as a focus of both sub- sistence and cultural identity. In cooperation with the Smithsonian Institution, I?upiaq Eskimo elders are interpreting the cultural and spiritual dimensions of whaling artifacts collected during the late nineteenth century, including material gathered by the Interna- tional Polar Expedition to Point Barrow, Alaska (1881? 1883). These artistic objects? hunting and boat equipment, regalia for whaling ceremonies, and charms owned by whale boat captains (umialgich)? were acquired during decades of rapid cultural change brought about by interaction with New England whalers, traders, and Presbyterian mis- sionaries. Nonetheless, the social values and spiritual concepts that they express have survived and are carried forward in contemporary whaling. Current research and ex- hibitions benefi t from both I?upiaq expertise and a rich ethnohistorical literature from Barrow and other northern communities. INTRODUCTION In the northern Bering Sea and Arctic Ocean, a 2,000-year tradition of Alaska Native whaling continues to the present day (Brewster, 2004; Freeman et al., 1998; McCartney, 1995, 2003). The spring bowhead hunt in particular? and the preparations and celebrations that surround it? are a focus of cultural iden- tity and survival (Worl, 1980). There are eight contemporary I?upiaq whaling villages: Nuiqsut, Barrow, Wainwright, Point Hope, Kivalina, Kaktovik, Wales, and Little Diomede. Two Yupik whaling communities, Gambell and Savoonga, are located on St. Lawrence Island. In cooperation with the Arctic Studies Center (Department of Anthropol- ogy, National Museum of Natural History, Smithsonian Institution), I?upiaq Eskimo community members are reexamining this ancient hunting heritage through the study of traditional whaling equipment in the collections of the Smithsonian Institution. This project is in part a legacy of the fi rst International Polar Year. During the U.S. government? sponsored International Polar Expe- dition to Point Barrow, Alaska (1881? 1883), commander Lt. Patrick Henry Ray and naturalist John Murdoch purchased more than 1,100 items from lo- cal I?upiaq residents including a wide variety of clothing, tools, and hunting Aron L. Crowell, Alaska Director, Arctic Studies Center, National Museum of Natural History, Smithsonian Institution, 121 W. 7th Avenue, Anchorage, AK 99501, USA (crowella@si.edu). Accepted 9 May 2008. The Art of I?upiaq Whaling: Elders? Interpretations of International Polar Year Ethnological Collections Aron L. Crowell 09_Crowell_pg099-114_Poles.indd 9909_Crowell_pg099-114_Poles.indd 99 11/17/08 8:37:35 AM11/17/08 8:37:35 AM 100 SMITHSONIAN AT THE POLES / CROWELL weapons (cf. Burch, 2009, this volume; Fitzhugh, 1988; cf. Krupnik, 2009, this volume; Murdoch, 1892; Ray, 1885). Of special signifi cance for the present discussion is a group of about 40 objects related to bowhead whal- ing and whaling ceremonies. In 2002, a group of cultural advisors from Barrow? Ronald Brower Sr., Jane Brower, Kenneth Toovak, and Doreen Simmonds? visited Wash- ington, D.C., to examine some of the Murdoch? Ray materials, as well as other collections at the National Museum of Natural History (NMNH) and National Mu- seum of the American Indian (NMAI) (Figure 1). The lat- ter include objects from Barrow, Point Hope, Little Dio- mede, and Wales that were acquired in the late nineteenth century by Edward W. Nelson, Miner Bruce, George T. Emmons, J. Henry Turner, H. Richmond Marsh, and others. Additional contributions to the indigenous docu- mentation of these collections were made by Barrow elder and translator Martha Aiken, as well as Norton Sound region advisors Jacob Ahwinona (White Mountain) and Marie Saclamana (King Island/Nome). This project is one focus of the Arctic Studies Center?s Sharing Knowledge program, which seeks to document in- digenous oral histories and contemporary knowledge about objects in the Smithsonian?s Alaskan collections (Crowell and Oozevaseuk, 2006). Outcomes include the Sharing Knowledge website (http://alaska.si.edu) and a large col- laborative exhibition on Alaska Native cultures that will open at the Anchorage Museum in 2010. In cooperation with the University of Alaska, Fairbanks, the I?upiat Heri- tage Center in Barrow produced its own community-based exhibition in 2005, The People of Whaling (http://www.uaf .edu/museum/exhibit/galleries/whaling/ index.html). Its title places whaling at the core of cultural identity, underlined by the exhibition?s theme statement, which reads, ?Whal- ing is central to our lives. We continue to teach our youth to show respect for the whale and to share the harvest with the whole community.? CONTACT AND CHANGE IN I?UPIAQ WHALING The Smithsonian collections were acquired during de- cades of rapid cultural change. The American commercial whaling fl eet came to the Bering Sea in 1848 and the Bar- row area in 1854 (Bockstoce, 1986). During the 1880s, FIGURE 1. Left to right: Kenneth Toovak, Jane Brower, and Ron Brower Sr. at the National Museum of the American Indian, 2002. (Photo by Aron Crowell) 09_Crowell_pg099-114_Poles.indd 10009_Crowell_pg099-114_Poles.indd 100 11/17/08 8:37:35 AM11/17/08 8:37:35 AM THE ART OF I?UPIAQ WHALING 101 the whaling industry shifted its focus to shore-based oper- ations that employed many I?upiaq residents and greatly increased the direct infl uences of nonnative culture and economy (Cassel, 2003). Although the fi rst shore sta- tion at Barrow was established in 1884, the year after the Point Barrow expedition ended, Murdoch noted that Na- tive whalers had already acquired breech-loading guns for hunting caribou and ?plenty of the most improved modern whaling gear? through their contacts with the commer- cial fl eet (1892:53). The new weapons were bomb-loaded harpoons (called ?darting guns?) and shoulder guns that rapidly replaced traditional stone-tipped harpoons and lances. Local residents were also ?now rich in iron, civi- lized tools, canvas, and wreck wood, and in this respect their condition is improved? (Murdoch, 1892:53). In other ways, Murdoch observed, the community was in decline through the ?unmitigated evil? of the alcohol trade, social disruption caused by the American sailors, and the effects of introduced diseases. Indigenous population losses from infl uenza and other epidemics were severe, and food short- ages came about as caribou dwindled and whaling compa- nies depleted whale and walrus herds. I?upiaq willingness to sell or barter traditional items of material culture to collectors may have been related to these transformations in culture and living conditions (Fitzhugh, 1988). Beginning in 1890, Presbyterian missionaries actively sought to suppress shamanism, whaling rituals, and hunt- ing ceremonies, along with the spiritual concepts that underlay these practices (Figure 2). I?upiaq qargit, or cer- emonial houses, closed down under missionary pressure, FIGURE 2. Men dancing in a qargi (ceremonial house) at Wales before the whale hunt, 1901. (Photograph by Suzanne R. Bernardi, Anchorage Museum archives B96.9.06) 09_Crowell_pg099-114_Poles.indd 10109_Crowell_pg099-114_Poles.indd 101 11/17/08 8:37:39 AM11/17/08 8:37:39 AM 102 SMITHSONIAN AT THE POLES / CROWELL ending the ceremonies that had taken place inside (Larson, 1995). The last I?upiaq qargi at Point Hope became inac- tive in 1910, although social distinctions based on family membership in the former ceremonial houses continued to be important (Larson, 2003; VanStone, 1962; Burch, 1981). I?upiaq whaling captains, or umialgich, retained their pivotal social and economic roles, as did their wives (Bodenhorn, 1990), but the once-extensive spiritual duties of these positions declined. Despite the effects of Western contact, substantial con- tinuities are evident between arctic whaling communities of the past and present (Anungazuk, 2003; Braund and Moorehead, 1995). The bowhead harvest is substantial, with a current allotment from the International Whaling Commission (IWC) of 56 landed whales per year, to be divided among the 10 Alaskan villages. The IWC quota is based on historic per capita averages of the subsistence harvest between 1910 and 1969 (Braund and Associates, 2007). Whaling captains and their kin-based crews and wives work throughout the year to prepare for whaling, carry out the hunt, process the catch, and distribute the meat and blubber. Whaling boats with hand-built wooden frames and skin covers are still employed in some of the vil- lages, including Barrow. The Whale Festival or Nalukataq has survived and the Messenger Feast has been revived in modern form. Spiritual conceptions of whaling also per- sist. Ronald Brower Sr. said, Whalers respect their prey very highly . . . Whaling is a very important part of our life. In many ways, it?s part of our sa- cred beliefs. Everything that we?re doing in a year is dealing with whaling? some form of preparation, celebration, rites and ritu- als of whaling. 1 Elders? commentaries in Washington provide insight into connections between modern and traditional whal- ing, including the deep-seated cultural view that whales are sentient beings that respond to human ritual and re- spect by giving themselves to feed the community. THE ANNUAL CYCLE OF I?UPIAQ WHALING: PAST AND PRESENT Sources on late-nineteenth and early-twentieth-century I?upiaq whaling include primary observations (Murdoch, 1892; Nelson, 1899; Ostermann and Holtved, 1952; Ray, 1885; Simpson, 1875; Stef?nson 1919; Thornton, 1931), later anthropological and ethnohistorical reconstructions (Curtis, 1930; Rainey, 1947; Spencer, 1959; VanStone, 1962), and retrospective oral history and life stories (Brew- ster, 2004; Pulu et al., 1980). Most available information pertains to Barrow, Point Hope, and Wales. A brief synopsis of this diverse material, with comparisons to contemporary practices, is offered here as a foundation for the Smithson- ian discussions with elders. PREPARATIONS FOR WHALING In the traditional whaling pattern, I?upiaq crew- men worked through the winter and early spring in the qargit (ceremonial houses) owned by their captains, pre- paring gear for the coming hunt. Everything? harpoons, lances, fl oats, whaleboat (umiaq) frames? had to be newly made or scraped clean in the belief that no whale would approach a crew with old or dirty gear (Curtis, 1930:138; Rainey, 1947: 257? 258; Spencer, 1959: 332? 336). The cap- tain?s wife supervised the sewing of a new bearded seal or walrus hide cover for the umiaq; the women assigned to this task worked in an ice-block house adjacent to the qargi. Women made new parkas and boots for the hunters. Ritual equipment was prepared for the whaling captain?s wife, including a wooden bucket with ivory ornaments and chains (Figure 3). The umialik (whaling captain) con- sulted with shamans and advisors, seeking ritual advice and portents of the season. He cleared out his ice cellar, distributing any meat from the previous year, to make way for the ?parkas? or fl esh of new whales, which the whale spirits shed for human use (Rainey, 1947:259; Spencer, 1959: 335? 336). White beluga whales are often the fi rst to be seen in the open water leads of early spring; at Wales, these were considered to be the bowheads? scouts, sent ahead to see if the village was clean and ready (Curtis, 1930:152). The bowheads would soon follow, and it was time to clear a path for the boats across the sea ice to the water?s edge. Boat captains retrieved whaling charms that they had hid- den in secret caches and caves (Spencer, 1959:338? 340). When charms were placed in the umiaq, it became a liv- ing being. At Wales, it was said that on the night before the hunt, the boats walked out to sea using the posts of their racks as legs (Curtis, 1930:152). Before the umiaq was launched, the captain?s wife gave it a drink of water from her bucket (Rainey, 1947:257). She herself was iden- tifi ed with the whale, and at Point Hope the harpooner pretended to spear her before the crew set out (Rainey, 1947: 259). Women still sew new umiaq covers each season in the I?upiaq whaling communities, although the activity has moved to alternative locations; at Barrow, it takes place 09_Crowell_pg099-114_Poles.indd 10209_Crowell_pg099-114_Poles.indd 102 11/17/08 8:37:42 AM11/17/08 8:37:42 AM THE ART OF I?UPIAQ WHALING 103 inside the I?upiat Heritage Center. Clean, new gear and clothing are considered to be just as essential now as they were in the past, to show respect for the whales (Brewster, 2004; Bodenhorn, 1990). Ivory whale charms are carried onboard some umiat; in others, the traditional image of a bowhead is carved on the underside of the boat steerer?s seat (see discussions below). Ice cellars are cleaned each spring, to make a welcoming home for the whales? bodies. THE HUNT Traditionally, women could not sew during the hunt because the act of stitching or cutting might entangle or break the harpoon line. A whaling captain?s wife sat quietly in her house to mimic a docile whale that would be easier to catch. She would not stoop or go into an underground meat cellar, acts that could infl uence a wounded bowhead to go under the ice where it would be lost (Rainey, 1947: 259; Spencer, 1959: 337? 338). As hunters approached a whale, the harpooner raised his weapon from an oarlock-shaped rest in the bow, thrusting it at close quarters into the animal?s back. The whale dove with the harpoon head inside its body, drag- ging the attached line and sealskin fl oats. The harpooner refi tted his weapon with a new head, prepared to strike again when the whale resurfaced. Other boats joined in the hunt. The whale eventually lay exhausted on the sur- face where it was killed with stone-tipped lances (Mur- doch, 1892: 275? 276; Rainey, 1947: 257? 259) The captain?s wife greeted the whale at the edge of the ice. Singing and speaking a welcome, she poured fresh wa- ter on its snout from her ceremonial bucket (Curtis, 1930: 141; Osterman and Holtved, 1952:26; Spencer, 1969: 345; Stef?nsson, 1919:389). Yupik and Siberian whal- ing cultures shared this practice of quenching a whale?s assumed thirst for fresh water. I?upiat traditionally gave all sea mammals they killed a drink of fresh water, and all land animals a taste of seal or whale blubber (which was rubbed on their noses), in the belief that the crea- tures of land and sea craved these substances that were not available to them in life (Brower, 1943:16; Rainey, 1947: 267; Spencer, 1969: 272; Stef?nsson, 1919: 389; Van Valin, 1941: 199). The contemporary spring whale hunt follows much the same course as in times past, with hunting crews camped at the ice edge by the beginning of May ( Brewster, 2004: 131? 163). Most crews use skin-covered umiat (propelled by paddles rather than motors during the hunt itself) to approach whales because the traditional hulls are much quieter in the water than wood or metal skiffs. The har- pooner with his darting gun and gunner with his shoulder gun ride in the bow, ready to strike with the aim of an immediate kill. Technological innovations go beyond the adoption of these now antique weapons. Whalers use snow machines and sleds to pull their boats and gear to camp, as well as modern communications equipment to enhance the logistics and safety of the hunt. They are aided by VHF radios, walkie-talkies, GPS units, satellite phones, and In- ternet forecasts of ice and weather conditions. At Barrow in 1997, when shorefast ice broke away and set 142 whal- ers adrift in heavy snow and fog, helicopters were guided to the rescue using GPS coordinates transmitted by radio from the men on the ice (George et al., 2004). Some of the older hunting prescriptions and prohi- bitions are retained, while others have faded. Today, as in the past, quiet is maintained in the whale camps because of the animals? sensitive hearing. On the other hand, cooking food on the ice is common practice now but banned under traditional norms (Rainey, 1947:259; Spencer, 1959:337). FIGURE 3. Umialik sitting beneath whaling charms, including women?s pails with ivory chains, Wales, 1901. (Photograph by Suzanne R. Bernardi, Anchorage Museum archives B96.9.05) 09_Crowell_pg099-114_Poles.indd 10309_Crowell_pg099-114_Poles.indd 103 11/17/08 8:37:43 AM11/17/08 8:37:43 AM 104 SMITHSONIAN AT THE POLES / CROWELL Captains? wives try to remain peaceful and quiet to infl u- ence the whale?s decision to give itself (Bodenhorn, 1990). Although she may no longer provide the customary drink of water to the whale, the captain?s wife and her husband act as ?good hosts? to the animal by sharing its meat with others during the Nalukataq celebration and at Christmas and Thanksgiving. CEREMONIES AND CELEBRATIONS The principal I?upiaq whaling ceremony, Nalukataq, represents an unbroken tradition that extends from pre- contact times to the present (Brower, 1943: 61? 63; Curtis, 1930: 135? 160; Larson, 2003; Murdoch, 1892: 272? 275; Spencer, 1969: 332? 353; Rainey, 1947:262). Nalukataq follows the whaling season and is celebrated outdoors, so it was minimally affected by the decline of the ceremonial houses. Each successful umialik provides a feast of whale meat and maktak (skin and blubber) to the entire village, an act that brings great prestige. Whaleboats tipped on their sides, with the fl ags of each crew fl ying, surround the outdoor space. Feasting is followed by dancing, singing, and competitive games, including the ?blanket toss? that gives the festival its name. During Nalukataq everyone re- ceives new boots and parka covers. The Apugauti feast marks the last time a successful whaling captain brings his boat back to shore at the end of the whaling season. It is a celebration of the boat?s re- turn. The captain raises his fl ag and everyone is invited to eat mikigaq (fermented whale meat), whale tongue, and maktak. Wild goose soup is also served. Kivgiq, the Messenger Feast, is a winter dance and gift? giving festival that was once widespread across north- west Alaska and in the Yup?ik regions of Norton Sound, the Yukon? Kuskokwim Delta, and Nunivak Island ( Bodfi sh, 1991: 23? 24; Burch, 2005: 172? 180; Curtis, 1930: 146? 147, 168? 177, 213? 214; Kingston, 1999; Lantis, 1947: 67? 73; Nelson, 1899: 361? 363; Oquilluk, 1973: 149? 150; Ostermann and Holtved, 1952: 103? 112; Spencer, 1957: 210? 228). I?upiaq Messenger Feasts ended in the early years of the twentieth century, but North Slope Borough Mayor George Ahmaogak Sr. helped to revive the event at Barrow in 1988. Barrow?s biennial celebration in February now brings visitors and dance groups from across Alaska, Russia, Canada, and Greenland. Before a traditional Messenger Feast, leading men of a village (usually the whaling captains) sent messengers to the leaders of another community to invite them and their relatives to fi ve days of ceremonies. At Utqiag? vik (Barrow), guests came to the qargi to view the great pile of gifts that they would receive, including sealskins fi lled with oil, weapons, sleds, and kayaks. While not specifi - cally a whaling ceremony, Kivgiq was an expression of the whaling-based coastal economy and of the social domi- nance of the umialgich. Before the disappearance of the ceremonial houses, fall and winter were a time for other whaling and hunting ceremonies. At Barrow the whaling season was followed by a feast and dance in the qargi, when men wore several types of masks. Few details were recorded or remembered about these dances, at least at Barrow. Murdoch pur- chased a dozen masks that were used in the qargi ceremo- nies, but had no opportunity to learn about their use. At Point Hope, hunting ceremonies were held each winter in the ceremonial houses until 1910 and elders? descriptions of these ceremonies were recorded by Froelich Rainey in 1940 (Rainey, 1947). Figures of whales, seals, polar bears, caribou, walrus, and birds were carved, hung in the qargi, and fed as part of the ritual. A mask with inset ivory eyes was hung above the oil lamp and the whaling captains vied with each other to steal it unobserved. It would be hidden in the victor?s cache and used in the spring as a whaling charm. SELECTED OBJECT DISCUSSIONS HARPOON REST (NAULIGAQAG ? VIK) Harpoon rests were fastened inside the bows of whal- ing boats, as a place for the harpooner to support his weapon (Murdoch, 1892: 341? 343; Nelson, 1899:226; Spencer, 1959: 342? 343). These implements are often dec- orated with whale imagery and were probably regarded as hunting charms. At Wales in 1927, a wooden harpoon rest was found in one umialik?s old cache of whaling talismans (Curtis, 1930:138). A walrus ivory harpoon rest from the Murdoch? Ray collection (Figure 4) is etched with images of bowhead fl ukes and each prong depicts a whale?s head and fore body (Murdoch, 1892: fi g. 348). Ronald Brower Sr. noted that on each prong the whale?s back is inset with a blue bead at the center of an inscribed ?X.? This, he explained, is the location of the whale?s life force, and the place where the harpooner aimed. Brower added that when blue is present on a hunting implement it is ?part of the weapon? and not just added for beauty. 2 Blue, he said, gives us the relationship to our spiritual beliefs, to the power of sixa? meaning ?sky?? who controls life. Sixa is something 09_Crowell_pg099-114_Poles.indd 10409_Crowell_pg099-114_Poles.indd 104 11/17/08 8:37:44 AM11/17/08 8:37:44 AM THE ART OF I?UPIAQ WHALING 105 that?s both fi nite and infi nite; you breathe it. When you look in the heavens you see blue. So blue became an important color that helped to bring a whale home. 3 Another harpoon rest from the village of Wales (Fig- ure 5), acquired by E. W. Nelson in 1881 (Nelson, 1899: Pl. LXXVIII? 37), is made from two pieces of walrus ivory that are pinned together with ivory pegs. Tihmiaqpat ( ?giant eagles?) in the act of catching whales are etched on the front and back, and animals with lifted paws? possibly polar bears? are carved on each side. Stories about giant eagles (or ?thunderbirds?) that preyed on whales, caribou, and people are found in the oral traditions of I?upiaq, Yup?ik, Chukchi, Koryak, St. Lawrence Island Yupik, Unangan, and other North Pacifi c peoples (e.g. Bogoras, 1904? 1909:328; Curtis, 1930: 168? 177; Ivanov, 1930: 501? 502; Jochelson, 1908: 661; Nelson, 1899: 445? 446, 486? 487). Despite their fearsome reputation, it was one of these birds, called the Eagle Mother, who is said to have taught I?upiaq people the dances and songs of the Messenger Feast (Kingston, 1999). Examining this harpoon rest, Jacob Ahwinona of White Mountain said, According to my grandpa, these birds, they?re up in the big mountains back there, way up high. When they go from there, they go out to the sea and pick those whales up, just like these ea- gles in the rivers pick salmon up [with their talons]. That?s right here, see? That bird is picking up that whale there, and then they bring them back to those high mountains. That?s where they nest. And when they bring those back, those bugs that grow there eat some of the leftovers from the bird?s nest. Those bugs that crawl there, my grandma said they?re as big as young seals. 4 Ahwinona reported that only a few years ago, when he was squirrel hunting at Penny River near Nome, a large shadow passed over the ground on a cloudless day, per- haps cast by one of the giant birds on its way out to the Bering Sea. FIGURE 4. Harpoon rest, Barrow, 1881? 1883, Murdoch? Ray col- lection. NMNH E089418. 30 cm tall. FIGURE 5. Harpoon rest, Wales, 1881, E. W. Nelson collection. NMNH E048169. 15 cm tall. 09_Crowell_pg099-114_Poles.indd 10509_Crowell_pg099-114_Poles.indd 105 11/17/08 8:37:45 AM11/17/08 8:37:45 AM 106 SMITHSONIAN AT THE POLES / CROWELL BOX FOR HARPOON BLADES (IKOIG ? VIK, ?STORAGE BOX?) A wooden container to hold and protect spare blades for the whaling harpoon was carried in the umiaq during hunting. Whaling captains cached the box after the whal- ing season, along with other hunting charms, amulets, and ritual objects (Bockstoce, 1977: 102; Curtis, 1930: 138? 139; Kaplan et al., 1984; Kaplan and Barsness, 1986:138; Murdoch, 1892: 247? 250; Nelson, 1899: 163, 439). Blade storage boxes were believed by I?upiaq whalers to infl u- ence hunting success (Nelson, 1899: 439) although exact conceptions about them were not historically recorded. The boxes are usually shaped like whales, but some repre- sent other animals such as polar bears and birds. Elders examined this whale-shaped box, collected by G. T. Emmons in about 1900 (Figure 6), as well as simi- lar examples in the Murdoch? Ray collection ( Murdoch, 1892: 246? 248). Its belly holds four triangular slate blades, secured beneath a wooden lid. The animal?s tail is shown with cut-off tips, an apparent reference to the traditional practice of cutting off the ends of the fl ukes or fl ippers and sending them to the captain?s wife to announce a suc- cessful hunt (Curtis, 1930: 140? 141; Rainey, 1947: 260; Spencer, 1959: 344). Today, as elders commented, the tips of a whale?s tail are still removed but for practical reasons, to reduce drag when towing the animal at sea. Brower likened the box to the ammunition cases that captains now carry in their boats to hold whale bombs, saying, From a spiritual sense, my observation is that many of our whalers still retain some of the old beliefs. But instead of a box like this, we now have a whale gun box, where we have our ammunition. This would be like an ammunition box in the old days. Now we have what we call the bomb box today, where we keep all of these same types of implements, used for the purpose of killing the whale. 5 Referring to the blade box, Kenneth Toovak said, ?They?re so powerful, some of these charms.? 6 He re- marked that they were probably owned by shamans and that Point Hope men possessed a powerful system of whal- ing ?medicine? in which boxes like these were employed. Brower concluded, This one is a box for the whales. The traditional beliefs deal with the spirit of the whale, and the spirit of the whale and the spirit of man are both intertwined. It is expected that whales gave themselves to the whalers. They not only are giving them- selves to the whalers, but to the captain?s wife, who has a ritual. Because that person maintained a clean household. Because the spirit of the whale is believed to be that of a girl. 7 Brower refers here to the identifi cation of the captain?s wife with the female spirit of the whale, both evoked by the artistic imagery of the blade box. Moreover, the whale gives its life not to the whaler but to his wife, in recog- nition of her skills, generosity, and observance of ritual (Bodenhorn, 1990). BOAT SEAT (Aqutim Iksivautaha, ?BOAT STEERER?S SEAT?) At Point Hope, wooden plaques carved with whale images were wedged inside the bow of the umiaq, mak- ing a small deck just in front of the harpooner. The whale fi gure was on the bottom side, facing downward and thus invisible. Froelich Rainey reported in 1940 that the har- pooner tapped the top of the platform while he sang a song that summoned hidden whales to the surface ( Lowenstein, 1993: 150). Talking about this example from the village of Wales (Figure 7), Barrow elders stated that it could be placed either in the bow of the boat (as at Point Hope) or in the stern as a seat for the boat steerer. In both places, the whale image would be on the bottom side. When used as a bow platform, Toovak said, the plaque would hold the coiled ak?unaaq (bearded sealskin line) that attaches to the whale harpoon. FIGURE 6. Whale-shaped box for holding harpoon blades, loca- tion unknown, 1900, G. T. Emmons collection. NMNH E204778. 45 cm long. 09_Crowell_pg099-114_Poles.indd 10609_Crowell_pg099-114_Poles.indd 106 11/17/08 8:37:48 AM11/17/08 8:37:48 AM THE ART OF I?UPIAQ WHALING 107 A carved whale seat/platform belongs to the umialik. Toovak said, ?Uvvakii ag? viqsiuqtinmakua umiag? iratih piqpagipiag? atag? uugait qutchiksuag? isuugait. Tavra tainna umialguruam marra sug? auttahi. ? (?And so it is whalers really do have respect for their boats and have high regard. These are a boat captain?s items.?) Brower said that the im- age of the whale is present in the boat as part of the ritual of whaling, and that its use is part of I?upiaq sacred beliefs. 8 The practice continues, as he noted in an earlier discussion. ?Some whale boats still have an ivory effi gy of the whale, tied on to the boat. Or they have an effi gy of the whale un- der the seat of the steersman in the rear of the boat.? 9 IVORY WHALING CHARMS (QAAGLI?IQ, ?CHARM,? OR AANG ? UAQ, ?AMULET?) Murdoch collected 21 whale fi gures made of walrus ivory, wood, and soapstone that he identifi ed as possible whaling charms, to be carried on the umiaq (Murdoch, 1892: 402? 405). Among these are three small carvings (3? 4? long) made of darkly stained walrus ivory (Figure 8). The two smaller whales are a matched male? female pair (the far left and far right fi gures, respectively) and the larger fi gure in the center is another female, as identifi ed by depictions of the external sex organs. Brower called this type of fi gure a qaagli?iq, 10 meaning a ?charm? that could attract animals but not compel them; it has the ?power of bringing.? This he contrasted with more potent fi gures called tuung?aq (shaman?s helping spirit), which were employed by shamans to control the animal?s spirit and which have ?the power of killing.? 11 He and other elders used the term aang? uaq (amulet) more ambiguously, as a synonym for both of the above. FIGURE 7. Whale plaque/seat for umiaq, Wales, purchased 1958. NMAI 226908.000. 42.5 cm wide. FIGURE 8. Ivory whaling charms, Barrow, 1881? 1883, Murdoch? Ray collection. Left to right: NMNH E089324, E089325, E089323. Largest 15 cm long. 09_Crowell_pg099-114_Poles.indd 10709_Crowell_pg099-114_Poles.indd 107 11/17/08 8:37:49 AM11/17/08 8:37:49 AM 108 SMITHSONIAN AT THE POLES / CROWELL SOAPSTONE FIGURE OF WHALE (AANG ? UAQ, ?AMULET,? OR TUUNG ? AQ, ?SHAMAN?S HELPING SPIRIT?) Another whale image, about 5? long, collected by Murdoch (Figure 9) appears to have been carved from the bottom of an old soapstone pot (Murdoch, 1892:404). In the discussion among elders, this fi gure was called both aang?uaq (amulet) and tuung?aq (shaman?s helping spirit). Observing what appeared to be blood that had been rubbed on the image, Brower suggested that, ?Something like this could probably be carried by the shaman, and he would add blood from the whale. And so he carries with him the life force of the whale.? Comparing this object with the ivory whaling charms (above), he said, ?One has more of a life force than the other. One has more strength, depending on the strength of the shaman. This one? it was used as an amulet and ensured that the whale would be caught.? 12 Returning later to the topic of shamanism, Brower said, Kenneth [Toovak] and I were talking earlier. When I was de- scribing those things that were used by shamans, we are reminded by our elders that that kind of life has passed. It is over. And it?s something that we did not inherit, because the life has changed. The traditional lifestyle? before Christianity set in? is gone. And so are the powers associated with that. Because today our people have accepted a new faith and live a different lifestyle, which does not require the old way of life in order to be successful. 13 HEADBAND (NIAQUG ? UN) A headband acquired at Barrow by H. Richmond Marsh in 1901 is made of bleached skin, animal teeth (caribou or Dall sheep incisors), red beads, and sinew thread (Figure 10). Murdoch reported that headbands made of Dall (mountain) sheepskin with dangling stone fi gures of whales were ?the badge of a whaleman,? worn by the umialik and harpooner for spring preparatory rites and during the hunt itself (Murdoch, 1892: 142). Some of these headbands were also decorated with mountain sheep teeth. Also at Barrow, John Simpson observed that head- bands made of caribou skin and hung with caribou teeth were worn ?only when engaged in whaling? (Simpson, 1875: 243). Murdoch was not able to acquire an example during the Point Barrow expedition because these articles were highly prized and rarely offered for sale. Barrow elders were not familiar with this type of headband, which is no longer worn. They suggested that it might be a woman?s ornament worn during a traditional ceremony to mark the autumn equinox ?when two stars appear? (aagruuk)? also referred to as the I?upiaq New Year? or else during the Messenger Feast. 14 BUCKET FOR GIVING DRINK TO WHALE (IMIQAG ? VIK) AND BUCKET HANDLE (IPU) Elders identifi ed a wooden bucket from Wales (Fig- ure 11) in the NMAI collection as one that may have been used by a whaling captain?s wife to provide fresh water to a newly caught whale. Pails for this purpose were also made of baleen. Toovak and Brower identifi ed the bucket?s various attachments. The ivory carvings on the rim (each is a qi?iyunaqsaun, ?ornament?) represent polar bear heads and a whale. Several ?hunter?s items? hang on a leather cord: a polar bear tooth, a walrus tooth, part of a har- poon head, ivory weights for a bolas to hunt birds, and an ivory plug for a sealskin fl oat. Two more fl oat plugs (puvuixutahit) are tied to the handle. Brower suggested that all of these ornaments are symbols of success in both hunting and political leadership. ?Communities respect the headman. That is indicated by making him gifts of this na- ture (indicates bucket), especially if he?s a very successful hunter.? 15 FIGURE 9. Stone whaling amulet, Barrow, 1881? 1883, Murdoch? Ray collection. NMNH E089557. 15 cm long. 09_Crowell_pg099-114_Poles.indd 10809_Crowell_pg099-114_Poles.indd 108 11/17/08 8:37:52 AM11/17/08 8:37:52 AM THE ART OF I?UPIAQ WHALING 109 An ivory handle with whale fi gures from Sledge Island (Figure 12) is from the same type of bucket. The water giv- ing ceremony, Brower said, was to help the whale ?move from the ocean to the land.? 16 Historically, these buckets served in other rituals. An umialik?s wife gave a drink to the whaleboat when it was launched, because the sealskin-covered umiaq was itself viewed as a kind of living sea mammal (Brower, 1943: 48; Rainey, 1947: 257; Spencer, 1959:334; Thornton, 1931:166? 167). At Point Hope, women raised their buck- ets to Alignuk, the Moon Man who controlled game. Peo- ple said that if the water in the woman?s pot was clear and clean, Alignuk would drop a whale into it, meaning that her husband would be successful in the spring hunt (Pulu et al., 1980: 15? 16; Rainey, 1947: 270? 271; Osterman and Holtved, 1952: 228). At Point Hope and Barrow, skilled craftsmen made new buckets each year for the whaling captains and their wives. These were bigger each time to show the umialik?s growing experience. Buckets were initiated with songs and ceremonies in the qargi (Rainey, 1947: 245; Spencer, 1959: 334). MAN?S DANCE GLOVES (ARGAAK, ?PAIR OF GLOVES?) A pair of Point Hope dance gloves collected in 1881 by E. W. Nelson (Figure 13) is made of tanned caribou skin and decorated with strings of red, white, and blue beads and with alder-dyed fringes at the wrists (Nelson, FIGURE 10. Headband with mountain sheep or caribou teeth, Barrow, 1901, H. Richmond Marsh. NMNH E209841. 25 cm across. FIGURE 11. Woman?s ceremonial bucket, Wales, purchased 1952. NMAI 218952.000. 23 cm tall. FIGURE 12. Handle for ceremonial bucket, Sledge Island, 1881, E. W. Nelson collection. NMNH E044690. 30 cm long. 09_Crowell_pg099-114_Poles.indd 10909_Crowell_pg099-114_Poles.indd 109 11/17/08 8:37:54 AM11/17/08 8:37:54 AM 110 SMITHSONIAN AT THE POLES / CROWELL 1899:38, Pl. XX? 1). A keeper string to go around the neck is ornamented with copper cylinders and blue beads. Ac- cording to Kenneth Toovak, ?Sometimes a man had a special Eskimo dance song, an original song . . . Atuutiqag? uurut taipkuagguuq atug? uuramihnik (?they have short songs that they sang?) . . . And he had pre- pared gloves for the special song that he made. He added that these dance songs were performed at the summer whaling festival (Nalukataq) and winter hunt- ing ceremonies, usually by whalers; at Point Hope songs would strictly be the property of different clans. Gloves were worn only during the dance, and then removed, a custom that continues to be observed in modern perfor- mances. 17 CHILD?S SUMMER BOOTS (PI?IG ? AK, ?PAIR OF BOOTS?) Toovak recognized a pair of short summer boots (Fig- ure 14) as the type made for children to wear during Nalu- kataq. The thin soles are made from young bearded sealskin that has been chewed to soften and crimp the toes, sides, and heels. Jane Brower identifi ed the uppers as bleached sealskin dyed with alder bark, while the upper trim and straps are of plain bleached skin. Ronald Brower said, In the old days, all the men, women, and children dressed in their fi nest clothes after the feast, when they were beginning to do the celebrations and dances. Everybody, after they had eaten, put on their fi nest clothes, including little children. 18 DISCUSSION Ethnological recording and collecting was assigned a lower priority than other scientifi c work by the leader- ship of the Point Barrow Expedition of 1881? 1883, and it was only possible to carry it out during brief periods when ongoing magnetic and meteorological observations could be set aside. Murdoch?s Ethnological Results of the Point Barrow Expedition (1892), for all of its gaps and fl aws (especially with regard to social and ceremonial life), re- fl ects an extraordinary effort to overcome the limitations of time and opportunity in the fi eld (cf. Burch, 2009, this volume). It is probably safe to say that the ultimate value of this effort was not anticipated at the time. Perhaps more evi- dent to members of the expedition was that the Point Bar- row I?upiat were at a cultural turning point, buffeted by the growing social and economic impacts of commercial whaling. Although Murdoch felt that they were ?essen- tially a conservative people? who remained independent and were not yet overwhelmed by change, he cataloged the new pressures on their society. In the decades after the expedition, these pressures increased with the advent of shore-based whaling stations, Presbyterian missions, food shortages, and epidemics. What was certainly not foreseeable in 1883 was that more than a century later the northernmost Alaskan vil- FIGURE 13. Man?s dance gloves, Point Hope, 1881, E. W. Nelson collection. NMNH E064271. 30 cm long. FIGURE 14. Child?s short summer boots for Nalakutaq, location unknown, 1931, donated by Victor J. Evans. NMNH E359020. 16 cm tall. 09_Crowell_pg099-114_Poles.indd 11009_Crowell_pg099-114_Poles.indd 110 11/17/08 8:37:57 AM11/17/08 8:37:57 AM THE ART OF I?UPIAQ WHALING 111 lages would still be whaling and that the practice would remain a vital center point of the culture and way of life. Due to this surprising stability, whaling objects from the late nineteenth century remain as culturally legible sign- posts within a continuing tradition. Some nineteenth- century types are still in current use, such as carved ivory hunting charms. Others are interpretable within a per- sistent conceptual frame? the reciprocal relationship be- tween whales and people, with its obligations of ritual and respect. Also impossible to imagine in 1883 would have been the new uses of museum collections, especially the current strong emphasis on making them accessible for Alaska Native interpretation, cultural education, and community- based exhibitions (Crowell, 2004; Clifford, 2004; Fienup- Riordan, 1996, 2005). The collaborative work presented here is only preliminary. Limited by time and resources, only a few I?upiaq elders have so far been able to view the Smithsonian materials and only a small part of the to- tal Murdoch? Ray collection has been surveyed. I?upiaq consultants pointed out the necessity of involving elders from all of the whaling communities, so that each could comment on material from his or her village in greater depth. The opportunity for this will come as the objects discussed in this paper and more than 600 others from all of Alaska?s indigenous cultural regions are brought to An- chorage for exhibition in 2010. The Arctic Studies Center gallery in Anchorage will be designed not only for display but also as a research center where community members can remove every object from its case for study and dis- cussion. The Smithsonian collections, including the large number gathered as part of the fi rst International Polar Year, represent a scientifi c, cultural, and historical legacy that will continue to yield new meanings. ACKNOWLEDGMENTS My sincerest appreciation to the following individuals who shared their expert knowledge about whaling, I?upiaq culture, and the Smithsonian collections: Jacob Ahwinona (Kawerak Elders? Advisory Committee), Martha Aiken (North Slope Borough School District), Ronald Brower Sr (University of Alaska, Fairbanks)., Jane Brower (I?upiat Heritage Center), Kenneth Toovak ( Barrow Arctic Science Consortium), Marie Saclamana (Nome Public Schools), and Doreen Simmonds (I?upiat History, Language, and Culture Commission). Deborah Hull-Walski (National Museum of Natural History) and Patricia Nietfeld (Na- tional Museum of the American Indian) and their staffs assisted with the collections consultations in Washing- ton; coordinating at the Alaska end were Terry Dickey, Wanda Chin, and Karen Brewster of the University of Alaska Museum. Dawn Biddison (Arctic Studies Center, National Museum of Natural History) provided support- ing research and edited the transcripts and translations of the Washington consultations. Project funding and sup- port from the Rasmuson Foundation, National Park Ser- vice Shared Beringian Heritage Program, University of Alaska Museum, and Smithsonian Institution is gratefully acknowledged. Thanks to Igor Krupnik (Arctic Studies Center, National Museum of Natural History) and Karen Brewster (University of Alaska, Fairbanks) for reviews that helped to improve this paper. NOTES 1. Alaska Collections/Sharing Knowledge Project, Tape 34A:096? 112. 2. Alaska Collections/Sharing Knowledge Project, Tape 30A: 413? 430. 3. Alaska Collections/Sharing Knowledge Project, Tape 29A: 158? 172. 4. Alaska Collections/Sharing Knowledge Project, Tape 20B: 412? 434. 5. Alaska Collections/Sharing Knowledge Project, Tape 29A: 052? 059. 6. Alaska Collections/Sharing Knowledge Project, Tape 29A: 135. 7. Alaska Collections/Sharing Knowledge Project, Tape 29A: 152? 170. 8. Alaska Collections/Sharing Knowledge Project, Tape 34A: 031? 116. 9. Alaska Collections/Sharing Knowledge Project, Tape 29A: 063. 10. Alaska Collections/Sharing Knowledge Project, Tape 29A: 378. 11. Alaska Collections/Sharing Knowledge Project, Tape 31A: 166. 12. Alaska Collections/Sharing Knowledge Project, Tape 31A: 166? 220. 13. Alaska Collections/Sharing Knowledge Project, Tape 32A: 148? 159. 14. Alaska Collections/Sharing Knowledge Project, Tape 28A: 221? 311. 15. Alaska Collections/Sharing Knowledge Project, Tape 34A: 241? 334. 16. Alaska Collections/Sharing Knowledge Project, Tape 29A: 217? 259. 17. Alaska Collections/Sharing Knowledge Project, Tape 28A: 003? 037. 18. Alaska Collections/Sharing Knowledge Project, Tape 27A: 398? 436. LITERATURE CITED Anungazuk, H. O. 2003. ?Whaling: Indigenous Ways to the Present.? In Indigenous Ways to the Present: Native Whaling in the Western Arctic, ed. A. P. McCartney, pp. 427? 432. Salt Lake City: Univer- sity of Utah Press. Bockstoce, J. R. 1977. Eskimos of Northwest Alaska in the Early Nine- teenth Century: Based on the Beechey and Belcher Collections and 09_Crowell_pg099-114_Poles.indd 11109_Crowell_pg099-114_Poles.indd 111 11/17/08 8:38:01 AM11/17/08 8:38:01 AM 112 SMITHSONIAN AT THE POLES / CROWELL Records Compiled during the Voyage of H.M.S. Blossom to North- west Alaska in 1826 and 1827, ed. T. K. Penniman. University of Oxford, Pitt Rivers Museum Monograph Series No. 1. Oxford, U.K.: Oxprint Limited. ???. 1986. Whales, Ice, and Men: The History of Whaling in the Western Arctic. Seattle: University of Washington Press. Bodenhorn, Barbara. 1990. ?I?m Not the Great Hunter, My Wife Is.? I?upiat and Anthropological Models of Gender. ?tudes/Inuit/ Studies, 14(1? 2): 55? 74. Bodfi sh, W., Sr. 1991. Kusiq: An Eskimo Life History from the Arctic Coast of Alaska, ed. W. Schneider, L. K. Okakok, and J. M. Nageak. Fairbanks: University of Alaska Press. Bogoras, W. 1904? 1909. ?The Chukchee.? In The Jesup North Pacifi c Expedition, vol. 7, ed. F. Boas. New York: A. Stechert. Braund, S. R., and Associates. 2007. Quantifi cation of Subsistence and Cultural Need for Bowhead Whales by Alaska Eskimos. Prepared for the Alaska Eskimo Whaling Commission, Barrow. http:// www .iwcoffi ce.org/ _documents/ commission/ IWC59docs/ 59? ASW6.pdf (accessed 5 August 2007). Braund, S. R., and E. L. Moorehead. 1995. ?Contemporary Alaska Eskimo Bowhead Whaling Villages.? In Hunting the Largest Ani- mals: Native Whaling in the Western Arctic and Subarctic, ed. A. P. McCartney, pp. 253? 280. Edmonton: Canadian Circumpolar Insti- tute, University of Alberta. Brewster, K., ed. 2004. The Whales, They Give Themselves: Conver- sations with Harry Brower, Sr. Fairbanks: University of Alaska Press. Brower, C. 1943. Fifty Years Below Zero: A Lifetime of Adventure in the Far North, 4th ed. New York: Dodd, Mead, and Company. Burch, Ernest S., Jr. 1981. The Traditional Eskimo Hunters of Point Hope, Alaska: 1800? 1875. Barrow, Alaska: North Slope Borough. ???. 2005. Alliance and Confl ict: The World System of the I?upiaq Eskimos. Lincoln: University of Nebraska Press. ???. 2009. ?Smithsonian Contributions to Alaskan Ethnography: The IPY Expedition to Barrow, 1881? 1883.? In Smithsonian at the Poles: Contributions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 89? 98. Washington, D.C.: Smithsonian Institution Scholarly Press. Cassell, M. S. 2003. ?Eskimo Laborers: John Kelly?s Commercial Shore Whaling Station, Point Belcher, Alaska, 1891? 1892.? In Indigenous Ways to the Present: Native Whaling in the Western Arctic, ed. A. P. McCartney, pp. 387? 426. Salt Lake City: University of Utah Press. Clifford, J. 2004. Looking Several Ways: Anthropology and Native Heri- tage in Alaska. Current Anthropology, 45(1): 5? 30. Crowell, A. L. 2004. Terms of Engagement: The Collaborative Represen- tation of Alutiiq Identity. ?tudes/Inuit/Studies, 28(1): 9? 35. Crowell, A. L., and E. Oozevaseuk, 2006. The St. Lawrence Island Fam- ine and Epidemic, 1878? 1880: A Yupik Narrative in Cultural and Historical Context. Arctic Anthropology, 43(1): 1? 19. Curtis, E. S. 1930. The North American Indian, vol. 20, ed. F. W. Hodge. New York: Johnson Reprint Company. [Reprinted, 1970.] Fienup- Riordan, A. 1996. The Living Tradition of Yup?ik Masks: Aga- yuliyararput, Our Way of Making Prayer. Seattle: University of Washington Press. ???. 2005. Yup?ik Elders at the Ethnologisches Museum Berlin: Fieldwork Turned on Its Head. Trans. M. Meade, S. L?hrmann, A. Karlson, and A. Pauls. Seattle: University of Washington Press and Calista Elders Council. Fitzhugh, W. W., ed. 1988. ?Introduction.? In Ethnological Results of the Point Barrow Expedition, by John Murdoch, pp. xiii? xlix [orig. published 1892]. Washington, D.C.: Smithsonian Institution Press. Freeman, M. M. R., L. Bogoslovskaya, R. A. Caulfi eld, I. Egede, I. I. Krupnik, and M. G. Stevenson. 1998. Inuit, Whaling, and Sustain- ability. Walnut Creek, Calif.: AltaMira Press. George, J. C., H. P. Huntington, K. Brewster, H. Eicken, D. W. Norton, and R. Glenn. 2004. Observations on Shorefast Ice Dynamics in Arctic Alaska and the Responses of the I?upiat Hunting Commu- nity. ARCTIC, 7(4): 363? 374. Ivanov, S. V. 1930. ?Aleut Hunting Headgear and Its Ornamentation.? In Proceedings of the Twenty- third International Congress of Ameri- canists, New York, 1928, pp. 477? 504. Jochelson, W. 1908. ?The Koryak.? In The Jesup North Pacifi c Expedi- tion, vol. 6, ed. F. Boas. New York: A. Stechert. Kaplan, S. A., and K. J. Barsness. 1986. Raven?s Journey: The World of Alaska?s Native People. Philadelphia: University Museum, Univer- sity of Pennsylvania. Kaplan, S. A., R. H. Jordan, and G. W. Sheehan. 1984. An Eskimo Whal- ing Outfi t from Sledge Island, Alaska. Expedition, 26(2): 16? 23. Kingston, D. M. 1999. Returning: Twentieth- Century Performances of the King Island Wolf Dance. Ph.D. diss., University of Alaska, Fair- banks. Krupnik, Igor. 2009. ??The Way We See It Coming?: Building the Leg- acy of Indigenous Observations in IPY 2007? 2008.? In Smithso- nian at the Poles: Contributions to International Polar Year Sci- ence, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 129? 142. Washington, D.C.: Smithsonian Institution Scholarly Press. Lantis, M. 1947. Alaskan Eskimo Ceremonialism. American Ethnologi- cal Society, Monograph 11. New York: J. J. Augustin. Larson, M. A. 1995. ?And Then There Were None: The ?Disappear- ance? of the Qargi in Northern Alaska.? In Hunting the Largest Animals: Native Whaling in the Western Arctic and Subarctic, ed. A. P. McCartney, pp. 207? 220. Edmonton: Canadian Circumpolar Institute, University of Alberta. ???. 2003. ?Festival and Tradition: The Whaling Festival at Point Hope.? In Indigenous Ways to the Present: Native Whaling in the Western Arctic, ed. A. P. McCartney, pp. 341? 356. Salt Lake City: University of Utah Press. Lowenstein, T. 1993. Ancient Land: Sacred Whale. The Inuit Hunt and Its Rituals. New York: Farrar, Straus and Giroux. McCartney, A. P., ed. 1995. Hunting the Largest Animals: Native Whal- ing in the Western Arctic and Subarctic. Edmonton: Canadian Cir- cumpolar Institute, University of Alberta. ???, ed. 2003. Indigenous Ways to the Present: Native Whaling in the Western Arctic. Salt Lake City: University of Utah Press. Murdoch, J. 1892. Ethnological Results of the Point Barrow Expedition. Ninth Annual Report of the Bureau of American Ethnology 1887? 88, pp. 3? 441.Washington, D.C.: Government Printing Offi ce. Nelson, E. W. 1899. The Eskimo about Bering Strait. Eighteenth Annual Report of the Bureau of American Ethnology, 1896? 97, pp. 3? 518. Washington, D.C.: Government Printing Offi ce. Oquilluk, W. A. 1973. People of Kauwerak: Legends of the Northern Eskimo. Anchorage: Alaska Methodist University. Ostermann, H., and E. Holtved, eds. 1952. The Alaskan Eskimos as De- scribed in the Posthumous Notes of Dr. Knud Rasmussen. Trans. W. E. Calvert. Report of the Fifth Thule Expedition, 1921? 1924, Volume 10(3). Copenhagen: Nordisk. Pulu, T. I. (Qipuk), R. Ramoth- Sampson (Tatqavin), and A. Newlin (Ipi- ilik). 1980. Whaling: A Way of Life (Agvigich Iglauninat Nigin- mun). Anchorage: National Bilingual Materials Development Cen- ter, Rural Education, University of Alaska. Rainey, F. G. 1947. The Whale Hunters of Tigara. Anthropologi- cal Papers of the American Museum of Natural History, 41(2): 231? 283. Ray, P. H. 1885. ?Ethnographic Sketch of the Natives of Point Barrow.? In Report of the International Polar Expedition to Point Barrow, Alaska, in Response to the Resolution of the House of Representa- tives of December 11, 1884, Part III, pp. 37? 60. Washington, D.C.: Government Printing Offi ce. 09_Crowell_pg099-114_Poles.indd 11209_Crowell_pg099-114_Poles.indd 112 11/17/08 8:38:01 AM11/17/08 8:38:01 AM THE ART OF I?UPIAQ WHALING 113 Simpson, J. 1875. ?Observations on the Western Eskimo and the Coun- try They Inhabit, from Notes Taken During Two Years at Point Barrow.? In Arctic Geography and Ethnology, pp. 233? 275. London: Royal Geographic Society. Spencer, R. F. 1959. The North Alaskan Eskimo: A Study in Ecology and Society. Bureau of American Ethnology Bulletin 171. Washington, D.C.: Smithsonian Institution Press. Stef?nson, V. 1919. The Stef?nson? Anderson Arctic Expedition of the American Museum: Preliminary Ethnological Report. Anthropo- logical Papers of the American Museum of Natural History, 14(1). Thornton, H. R. 1931. Among the Eskimos of Wales, Alaska, 1890? 93, ed. N. S. Thornton and W. M. Thornton, Jr. Baltimore: Johns Hopkins Press. Van Valin, W. B. 1941. Eskimoland Speaks. Caldwell, Idaho: Caxton Printers. VanStone, J. 1962. Point Hope: An Eskimo Village in Transition. Seattle: University of Washington Press. Worl, R. 1980. ?The North Slope I?upiat Whaling Complex.? In Alaska Native Culture and History, ed. Y. Kotani and W. B. Workman, pp. 305? 321. Senri Ethnological Studies No. 4. Osaka, Japan: Na- tional Museum of Ethnology. 09_Crowell_pg099-114_Poles.indd 11309_Crowell_pg099-114_Poles.indd 113 11/17/08 8:38:02 AM11/17/08 8:38:02 AM 09_Crowell_pg099-114_Poles.indd 11409_Crowell_pg099-114_Poles.indd 114 11/17/08 8:38:02 AM11/17/08 8:38:02 AM ABSTRACT. As part of the First International Polar Year, the Smithsonian Institution established a meteorological and astronomical observatory at Ft. Chimo (Kuujjuaq) in Ungava Bay in 1881? 1883. Sent to man the post was the Smithsonian?s most prominent northern naturalist, Lucien Turner. Turner developed a close rapport with Inuit and Innu families from whom he acquired an extraordinary array of scientifi c specimens and ethno- logical materials. While intrepid and inspired, the work of the Smithsonian?s pioneering Arctic scientists refl ects the biases of western scientifi c tradition. Northern Native peoples were viewed as part of the arctic ecosystem to be observed, cataloged, and described. For the most part, the intellectual landscape of Innu and Inuit groups was overlooked and ignored. The Smithsonian collections are a powerful talisman for evoking knowledge, appreciation, and pride in Innu and Inuit heritage and serve as one point of departure for research during the Fourth IPY in 2007? 2008. Recognition that northern Natives have a mandate to participate in and inform northern research is an important change in the production of northern scientifi c research. INTRODUCTION This essay considers the changes in the practices of museum anthropology and archaeology at the Smithsonian Institution between the First IPY in 1882? 1883 and the current IPY of 2007? 2008. To know a place is to name it. The place we call the Arctic means different things to different people. A cultural construct defi ned by different eyes and different ways of knowing, it is both real and intangible. Archaeologist Robert McGhee (2007) calls it ?the last imaginary place.? It is only in the twentieth century that the technologies and the insatiable appetites of the developed world have been able to overcome environmental and logistical constraints to establish a permanent presence throughout the north. There are libraries and research institutes devoted to the complexity and variety of human experiences at high latitudes. For visitors, the Arctic is as much a cul- tural construct as it is a physical one, with perceptions repeatedly shaped and reshaped by time and circumstance. One has only to consider the transformation of Arctic landscapes from the fantastic fairy-tale visions? gothic cathedrals of ice? of the early-nineteenth- century explorers, subsequently morphed by suffering and danger into the grim Stephen Loring, Arctic Studies Center, Depart- ment of Anthropology, National Museum of Nat- ural History, Smithsonian Institution, P.O. Box 37012, MRC 112, Washington, DC 20013-7012, USA (lorings@si.edu). Accepted 9 May 2008. From Tent to Trading Post and Back Again: Smithsonian Anthropology in Nunavut, Nunavik, Nitassinan, and Nunatsiavut? The Changing IPY Agenda, 1882? 2007 Stephen Loring 10_Loring_pg115-128_Poles.indd 11510_Loring_pg115-128_Poles.indd 115 11/18/08 9:17:27 AM11/18/08 9:17:27 AM 116 SMITHSONIAN AT THE POLES / LORING and foreboding visions of the ubiquitous and relentless ice of the post-Franklin era (Figure 1), to the modern era with its coffee-table books of stunning photography of polar bears and vast unpopulated expanses (Loomis, 1977; 1986; Grant, 1998). But the Arctic is also a homeland, and has been for thousands of years. Arctic inhabitants have evolved a remarkable and practical adaptation to the cli- matic and ecological extremes of the northern polar world. Indigenous knowledge? based on observation and infer- ence and passed from generation to generation? forms an astute and perhaps surprisingly complex interpretation of the world inhabited by indigenous northern peoples. Yet with few exceptions (e.g., Rasmussen, 1929; Rink, 1875), visiting researchers and scientists have not learned the lan- guage of their hosts and thus have been denied much of the complexity of northern perceptions that has developed over generations by indigenous peoples. Historically, science in the north began as a hand- maiden of colonial enterprise. Having developed the tech- nology to transport them into (if not always out of) the polar regions, nineteenth-century explorers, with their passion for expanding scientifi c and geographic knowl- edge, began to collect information about the places they found themselves in. As was typical on many early and mid nineteenth-century Arctic voyages, with their winter quarters established, observatories were placed on the ice and rounds of tidal measurements, weather, and geophysi- cal observations began. Expedition accounts are fi lled with observations on the phenomenology of ice and cold, as navy explorers and scientists confronted the mysteries of Arctic life and returned with their collections of natural history specimens. Also in these accounts, are anecdotal and ethnohistorical passages that provide some of the fi rst descriptions of Native residents of the Arctic. In compari- son to more complex societies elsewhere, northern band- level societies, with their more modest material remains (the very antithesis of Euro-American values of domi- nance, competition and wealth) were perceived as being backward and marginal, literally frozen in time. Lacking a critical self-awareness the eyes of the European explor- ers had yet to take the true measure of the Natives of the eastern arctic who were frequently portrayed as quaint and childlike, devoid of the quarrelsome and bellicose at- titudes of some of their western and southern counterparts (Figure 2). Gradually the accumulation of geographical knowl- edge, dearly bought, began to make sense of the physi- cal mysteries of the arctic. Still little in the way of seri- ous attention to native cultures was afforded prior to the travels of Charles Francis Hall beginning in 1860 (Hall, 1864; Loomis, 1972). More visionary than scientist, Hall had been drawn to the arctic by the continuing fascina- tion with the fate of the lost Sir John Franklin Expedi- tion (1845) and the possibility that survivors might yet be living amongst the Inuit. Severely curtailed by fi nancial constraints Hall broke from the prevailing tradition of us- ing expedition ships frozen in the ice as base stations from which to launch sledge and small-boat voyages in favor of adopting Inuit modes of travel. Hall moved in with his Nunavut hosts, learned their language, and experienced their culture as an active participant. He was fortunate FIGURE 1. The terrible tragedy surrounding the loss of life during the U.S. North Polar Expedition (1879? 1881) following on the de- bacle of the British Northwest Passage Expedition under Sir John Franklin (1845? 1848) had soured public opinion in the United States on the benefi ts of polar exploration and transformed perceptions of the Arctic as a deadly and foreboding landscape. Editorial cartoon, Frank Leslie?s Illustrated Newspaper, 20 May 1882, William Dall papers, RU7073, SI Archives. (S. Loring photograph.) 10_Loring_pg115-128_Poles.indd 11610_Loring_pg115-128_Poles.indd 116 11/18/08 9:17:27 AM11/18/08 9:17:27 AM FROM TENT TO TRADING POST AND BACK AGAIN 117 in befriending an extraordinary Inuit couple, Ebierbing (?Joe?) and Tookoolito (?Hannah?) who provided Hall with an entr?e into Inuit society and served as his guides and guardians on all of Hall?s three arctic expeditions (Loomis, 1997). Hall?s receptivity to Inuit testimony and acceptance of the validity of Inuit knowledge about their history and their homeland both assured his own survival and the success of his expeditions and presaged (by more than half a century) the recognition of the validity and acuity of Inuit oral knowledge by subsequent Arctic scien- tists and travelers (Woodman, 1991) (Figure 3). In the eastern Arctic, it is not until nearly 20 years after Hall that the fl edgling discipline of Arctic anthro- pology emerged as a direct consequence of the fi rst Inter- national Polar Year with the arrival of Lucien Turner in Ungava in 1882? 1884 and Franz Boas in Baffi n Island in 1883? 1884 (Loring, 2001a; Cole and Muller-Wille, 1984). Neither Boas nor Turner was formally trained in anthropology. Turner?s fi rst interest and abiding passion was ornithology while Boas came to the eastern arctic as a geographer. Boas? trip to Baffi n Island was planned and partially sponsored by the German Polar Commission, which was then processing the data gathered from the Ger- man IPY station in Cumberland Sound (Barr, 1985). The resulting ethnographic monographs of both Boas (1885) and Turner (1894) were subsequently published by the Smithsonian Institution. These monographs have proved to be the emerging discipline of anthropology?s intellectual bedrock for research pertaining to the indigenous peoples of Baffi n Island and northern Quebec-Labrador, and these FIGURE 2. Illustrations accompanying William Edward Parry?s popular accounts of his search for the Northwest Passage (1819? 1834) depict the central Canadian Arctic Inuit as whimsical and childlike catered to a European perception of the polar region as a fantastic otherworldly place (Parry 1821, 1824). Detail from a Staffordshire ceramic plate, ?Arctic Scenery? ca. 1835. (Photograph by S. Loring of plate in author?s collection.) 10_Loring_pg115-128_Poles.indd 11710_Loring_pg115-128_Poles.indd 117 11/18/08 9:17:37 AM11/18/08 9:17:37 AM 118 SMITHSONIAN AT THE POLES / LORING works remain as some of the lasting triumphs of the fi rst IPY accomplishments. The scientifi c agendas of the International Polar Years, in 1882? 1883, 1932? 1933, and 1957? 1958 have all been concerned with addressing problems of meteorol- ogy, atmospheric science, and high-latitude geophysics. Yet, ironically, arguably the most lasting accomplishments of the American contribution to the First IPY? from the Point Barrow and Ungava stations? were the collections of natural history specimens (e.g., Dunbar, 1983) and eth- nographic materials that Smithsonian naturalists acquired around the fringes of their offi cial duties as weather ob- servers for the U.S. Army Signal Service (Murdoch, 1892; Turner, 1894; Nelson, 1899). Surprisingly, the volumes of atmospheric, oceanic, magnetic, and solar observations gathered at the dozen IPY 1882? 1883 stations did not yield the anticipated insights into global climatic and geophysi- cal regimes (Wood and Overland, 2006). Perhaps more signifi cant than the research results in the physical sciences was the establishment of a model for international scien- tifi c practice based on coordination and cooperation, and the recognition that the study of high latitudes (at both poles), as with the high seas, was an arena of international consequence and signifi cance. In company with the earth sciences and natural history, anthropology and archaeology were part of the expand- ing western economic, social, and intellectual hegemony of the nineteenth century. The construction of scientifi c knowledge about the world has, for the most part, pro- ceeded following well-defi ned western notions of logic and scientifi c explanation as the principle explanatory process for understanding the natural world and the place of human beings therein. Now, in the twenty-fi rst century, with much of the world?s cultural and biological diversity documented in at least a preliminary fashion, anthropol- ogy faces the challenge of recognizing, articulating, inter- preting, and preserving as broad a spectrum of humanity?s shared cultural diversity as possible. At the time of the fi rst IPY, many of the indigenous peoples of North America had been swept from their tra- ditional homelands. Secure in their northern redoubts of ice and stone, the Natives of the eastern Arctic had been spared much of the continental dislocation and genocide waged against indigenous communities in warmer climes. The inroads of European explorers, and later missionar- ies, whalers, and traders, had not signifi cantly impeded traditional Inuit subsistence practices nor had they in- truded far into their spiritual matters and beliefs. Under the leadership of Spencer Baird, the fi rst curator of the U.S. National Museum and the Institution?s second secre- tary (1878? 1887), and later, John Wesley Powell (Director of the B.A.E. 1879? 1902), Smithsonian anthropology? in the guise of the Bureau of American Ethnology? operated under a paradigm of salvage anthropology in the belief that Native American peoples were fated to gradually decline and disappear. Situated in the National Museum of Natural History (NMNH), Smithsonian anthropology had a decidedly materialist, collections-based orientation that was strongly infl uenced by biological sciences and Darwin?s evolutionary doctrines. Northern native cultures were seen as being somewhat uniquely divorced from his- tory due to their remote geography, and many theorists of the day considered them to be a cultural relic of Ice Age Paleolithic peoples, at the extremity of the scale in terms of human cultural variation. FIGURE 3. Thule ground-slate whaling harpoon endblade found by Charles Francis Hall?s Inuit companion Ebierbing, also known as ?Esquimaux Joe.? Historically, the role of Inuit guides and companions in the production of Arctic science was rarely acknowl- edged.SI-10153, Charles Francis Hall collection, NMNH. (S. Loring photograph) 10_Loring_pg115-128_Poles.indd 11810_Loring_pg115-128_Poles.indd 118 11/18/08 9:17:51 AM11/18/08 9:17:51 AM FROM TENT TO TRADING POST AND BACK AGAIN 119 The Smithsonian Institution?s previous interests in the eastern arctic? beginning with biological studies in Hudson?s Bay in the early 1860s, and support for the U.S. Eclipse Expedition to northern Labrador in 1860, as well as its close relationship with the Hudson?s Bay Company (Lindsay 1993)? provided a basis for a concerted study in the Ungava region. As part of the First International Polar Year 1882? 1883, the Smithsonian Institution partnered with the U.S. Signal Corps to establish meteorological and astronomical observatories at Point Barrow, Alaska, at Ft. Conger on Ellesmere Island, and at the Hudson?s Bay Company Post at Fort Chimo (Kuujjuaq) in Ungava Bay (Barr, 1985). Sent to man the post at Kuujjuaq was one of the Smithsonian?s most experienced northern naturalists, Lucien Turner, who had previously conducted important studies for the Smithsonian in the Aleutian Islands and Western Alaska (Turner, n.d.; 1886; Loring, 2001a). Lucien Turner (1848? 1909) was at the center of a small and talented band of young naturalists that were recruited by the Smithsonian?s second secretary, Spencer Baird. An accomplished ornithologist, linguist, and taxi- dermist Turner reveled in the opportunities for research and collecting in the North American arctic. Baird ar- ranged for Turner to be posted at the Hudson?s Bay Com- pany post at the mouth of the Koksoak River in Ungava Bay as a member of an IPY-sponsored meteorological ob- servatory for the U.S. Signal Corps. Turner?s arrival at Fort Chimo in 1882 was something of a surprise for the chief factor there, as news from the outside world only arrived once a year with the annual supply ship. Not easily rebuffed, Turner quickly estab- lished his observatory and took up the responsibilities of his post (Figure 4), both those pertaining to his IPY agenda and those dictated by his Smithsonian mandate. Although constrained by the demanding regime of his observation and recording obligations, Turner was able to develop a close rapport with Inuit and Innu families visiting the post, from whom he acquired an extraordinary array of scientifi c FIGURE 4. Lucien Turner at his observatory at the Hudson?s Bay Company Post at Fort Chimo (near present day Kuujjuak), 1881. (SI-6968) 10_Loring_pg115-128_Poles.indd 11910_Loring_pg115-128_Poles.indd 119 11/18/08 9:17:53 AM11/18/08 9:17:53 AM 120 SMITHSONIAN AT THE POLES / LORING specimens and ethnological materials (Turner, 1888, 1894; Loring 2001a) (see Figure 5). Unfortunately, no traces of a personal diary or let- ters survive from Turner?s time at Fort Chimo. Diligent archival research, at the Smithsonian and Hudson?s Bay Company archives, provide a few tantalizing clues to his rapport with the northern Natives he came into contact with (Loring, 2001a). However, for the most part these contacts are only dimly referred to as the source for knowledge about the local environment, animals (includ- ing mammals, birds, fi sh, and invertebrates), social rela- tions, and mythology. The contemporary ?intellectual landscape? of Innu and Inuit groups? the complex web of oral histories and observational knowledge pertaining to animals, weather, and the land? was largely overlooked and ignored by the IPY-era anthropologists, as their focus, stemming from the natural history approach of their mis- sions, was to categorize and describe the material culture of the people they encountered. Despite being confi ned by the intellectual framework of the day, Turner?s Ungava collections (as well as the collections made by Murdoch and Ray at Point Barrow in 1881? 1883) have become a powerful instrument for evoking knowledge, appreciation, and pride in Innu and Inuit heritage. They serve as a point FIGURE 5. Innu women and children visiting Lucien Turner at Fort Chimo, 1881. Photography was deemed an essential component of the work of the Smithsonian naturalists. As some of the earliest extent photographic images of northern Natives, they remain a prominent legacy of the fi rst IPY. (SI-6977) 10_Loring_pg115-128_Poles.indd 12010_Loring_pg115-128_Poles.indd 120 11/18/08 9:18:05 AM11/18/08 9:18:05 AM FROM TENT TO TRADING POST AND BACK AGAIN 121 of departure for research during this IPY in 2007? 2008, as explained below. Within the confi nes of their training and natural history proclivities, the Smithsonian?s Arctic natu- ralists had a demonstrated sensitivity to some aspects of native knowledge pertaining to the cultural and biological collections they acquired, though for the most part these are brief and anecdotal notations. Today, these notes, but more signifi cantly the objects themselves, are being reex- amined and reinterpreted by descendants from the com- munities from which the objects had come more than a century before. THE SHIFT IN INTELLECTUAL PARADIGM The recognition that arctic people have an intellec- tual, moral, and sociopolitical mandate to participate in and inform northern research marked a fundamental and dramatic shift in the practice of scientifi c research in the north. It did not arrive until the 1970s and more fi rmly, until the 1990s (Berger, 1977; Berkes, 1999; Nadasdy, 1999; Stevenson, 1996; Nicholas and Andrews, 1997) (see Figure 6). With the passage of the Native American Graves Pro- tection and Repatriation Act in 1990 (NAGPRA) and the National Museum of the American Indian Act (in 1996), the intellectual landscape as it pertains to the use and study of the Native American collections has been transformed into a museum anthropology that is more inclusive, more diverse, and contingent on Native participation and exper- tise (Crowell et al., 2001; Fienup-Riordan, 1996, 2005a, 2005b, 2007; Loring 1996, 2001b; Swidler et al., 1997; Thomas, 2000; Watkins, 2003; 2005; Zimmerman et al., 2003). It is in this context of cooperation and respect that the agendas of Smithsonian anthropology and IPY con- verge as specifi c information about objects in the museum collection are not only interpreted by knowledgeable el- ders and descendant community representatives but also serve as a touchstone or gateway to discussions about tra- ditional ecological and environmental knowledge. It thus seems appropriate, given the degree that human agency is implicated in climatic change, that anthropology for the fi rst time has been formally recognized as a goal of IPY polar science, under its new mandate: to investigate the cultural, historical, and social processes that shape the resilience and sustainability of circumpolar human societies, and to identify their unique contributions to global cul- tural diversity and citizenship. (ICSU/WHO 2007:13) SEEING AND BELIEVING: CHANGING PERSPECTIVES IN MUSEUM ANTHROPOLOGY At the Smithsonian Institution, the climate and phi- losophy of repatriation (Loring, 2001b; 2008), coupled with the moral and inspirational presence of the new Na- tional Museum of the American Indian (NMAI), has en- couraged the emergence of new practices and scholarship. FIGURE 6. ?We were real red-men in those days!? says Uneam Kat- shinak, a much revered Innu hunter, as he reminisces about being covered in blood while spearing caribou from a canoe as a boy. The continuity of traditional subsistence practices has anchored northern Native perceptions of their identity and their homeland belying the ?vanishing Indian? paradigm of nineteenth-century anthropology. (S. Loring photograph at the Tshikapisk-sponsored rendezvous at Kamestastin, Nitassinan, September 2000) 10_Loring_pg115-128_Poles.indd 12110_Loring_pg115-128_Poles.indd 121 11/18/08 9:18:22 AM11/18/08 9:18:22 AM 122 SMITHSONIAN AT THE POLES / LORING The new paradigm is evidenced by the signifi cant num- bers and variety of northern Native American and Inuit scholars, academics, artisans, and visitors who come to acknowledge, study, and appreciate the collections that were derived from their ancestors a century or more ago (Figure 7). The Smithsonian?s anthropology collections acquired during the First International Polar Year in Alaska and Nunavik had languished for almost a cen- tury, awaiting an appreciation of their signifi cance? fi rst as objects of art with The Far North exhibition at the National Gallery of Art (Collins et al., 1973) and then as symbols of cultural glory and scholarly wonder in a pair of precedent-setting exhibitions, Inua: Spirit World of the Bering Sea Eskimo in 1982 and Crossroads of Conti- nents in 1988 (Fitzhugh and Kaplan, 1982; Fitzhugh and Crowell, 1988). In opening the Smithsonian?s ?attic? and in returning to northern Natives an awareness of their material culture patrimony, the role of the museum has been radically transformed. Today, the Smithsonian col- lections at the NMNH and the NMAI form the largest holding of material culture pertaining to the heritage and history of North America?s indigenous peoples. With the passage of time and the miracle of conservation, these objects have undergone an extraordinary transformation from natural history specimens and anthropological curi- osities to become the foundation stones for contemporary community identity and heritage (Figure 8). The challenge of the next century is to accommodate this transforma- tion and incorporate new perspectives and knowledge. The future of anthropology in the museum will en- courage? and necessitate? new ways of thinking about the past that would require museum anthropologists and archaeologists surrender? or negotiate? their prerogative to interpret the past. Even more important, it is incumbent upon museum professionals to learn new ways of listening, new ways of recognizing the legitimacy of other voices, and other ways of knowing and accepting oral tradition as a valid interpretive tool (Tonkin, 1992). This sea change in museum anthropology and archaeology is inherent in the programs and initiatives that Smithsonian anthropology is conducting during the time of IPY 2007? 2008. A joint NMNH? NMAI exhibition project is bringing more than 500 Native Alaskan artifacts collected around the time of the First IPY to Anchorage, Alaska. The exhibit relies heavily on curatorial input from teams of Native Alaskan consultants affi rming the legitimacy of their perspective, knowledge, and link to their legacy and heritage. AN ENDANGERED PERSPECTIVE ON THE PAST With the passing of the Inumariit, the knowledgeable Inuit who lived in the country in the manner of their an- cestors, and the Tsheniu Mantushiu Kantuat, the old Innu hunters with special powers, so passes one of the last ves- tiges of the link to the intellectual landscape of hunting- foraging peoples. To anthropologists and even many indig- FIGURE 7. Community-orientated collection consultation and outreach has been a core concept of the Smithsonian?s Arctic Studies Center since its inception in 1991. Here George Williams, from the village of Mekoryuk on Nunivak Island, Alaska, points out construction details of a model kayak that had been collected by Henry B. Collins in 1927. Williams was part of a delegation of Nunivak elders and educators who visited the Smithsonian in March 1996. (S. Loring photograph) 10_Loring_pg115-128_Poles.indd 12210_Loring_pg115-128_Poles.indd 122 11/18/08 9:18:25 AM11/18/08 9:18:25 AM FROM TENT TO TRADING POST AND BACK AGAIN 123 enous people themselves, it becomes increasingly diffi cult to grasp the richness and complexity of the hunters? worlds as discerned through artifacts and museum exhibits. The insights and wisdom derived from centuries of intimate knowledge and experience on the land now exists as a much impoverished and fragmentary corpus. In Labrador, the 1918 infl uenza pandemic devastated Inuit communities and savaged Innu camps, killing off a generation of story- tellers and tribal elders and often leaving camps where only children and dogs survived. Soon, much of what remains of this specialized knowledge will only reside in libraries, museum collections, and in the clues that archaeologists might deduce. However, there is yet a great potential for the practice of archaeology in the north to be informed by the knowledge and perceptions of elders, the last people to be born in snow houses and tents and to have spent much of their lives as subsistence hunters and seamstresses. The sense of urgency is palpable, as the stock of elders? knowl- edge and perceptions is not renewable and will not be with us much longer. Subsistence strategies are being replaced by the market economy; communal social relations predicated on reciprocity and kinship are subordinated by government mandates and initiatives. The problem can be framed in a global perspective of diminishing ecological, biological, and cultural diversity. All of which begs the question: How important are ?old ways? and ?traditional subsistence practices? in the modern world? In Labrador, as elsewhere in the north, today?s Innu and Inuit youth are village-dwellers. Born in hospitals and brought up in isolated rural towns, northern young people have few opportunities to acquire country experiences and knowledge. There is a huge discrepancy between the past as experienced by their grandparents and the present. However well-meaning Canadian government policies may have been in advancing schooling, health care, and old-age pensions, the results have often been disastrous (Samson, 2003; Shkilnyk, 1985). Suicide rates in Labrador Innu communities are the highest recorded in the world and substance abuse is rampant (Samson et al., 1999). Fueled by chronic unemployment and inappropriate educational development, the impoverishment of village life? devoid of country values and skills with its concomitant pack- age of social, health and economic woes? is striking in comparison to ethnohistorical accounts which invariably describe the Innu as arrogant, self-suffi cient, ?tiresomely? independent, and proud (Cabot, 1920; Cooke, 1979). KNOWLEDGE REPATRIATION AND ARCHAEOLOGY In this situation, Smithsonian cultural research in the north becomes a component in the communities? re- sponse to the current heritage crisis and social dissolution. For the past decade, in close collaboration with Native elected offi cials, community leaders, and teachers, the Arc- tic Studies Center has pioneered community archaeology programs in Labrador with Inuit and Innu communities FIGURE 8. A drawing of the so-called magic doll collected by Luc- ien Turner in 1881 from Labrador Inuit visiting the Hudson?s Bay Company Post at Ft. Chimo (Kuujjuak) in Nunavik. Such unique specimens eloquently attest to the continuity of shamanistic practices in country-settings beyond the purview of missionaries and trad- ers. Transformed into museum specimens such objects still retain a tremendous potency to inspire and inform descendant community members of the people from whom they had been acquired. (Fig. 22 in Turner 1894, Smithsonian catalog number ET982, NMNH) 10_Loring_pg115-128_Poles.indd 12310_Loring_pg115-128_Poles.indd 123 11/18/08 9:18:27 AM11/18/08 9:18:27 AM 124 SMITHSONIAN AT THE POLES / LORING ( Loring, 1998; Loring and Rosenmeier, 2005). These pro- grams have sought to develop archaeological fi eld-schools that would provide Native youth with opportunities to ex- perience life in the country, acquire new job skills, and fos- ter self-esteem and pride in oneself and one?s heritage. This type of enterprise, generally called ?community archaeol- ogy,? especially as it is practiced in the north, is rooted in applied socially conscious advocacy anthropology. In addition to addressing the special scholarly questions that archaeologists commonly pose, community archaeology seeks additional goals that strive to empower and engage communities in the recognition and construction of their own heritage. In the north in general, and in Labrador specifi cally, community archaeology initiatives celebrate traditional values and share a research focus and practice that is responsible for creating and returning knowledge to communities (Lyons, 2007; Nicholas, 2006; Nicholas and Andrews, 1997), in a sense coming full circle since the years of the fi rst IPY. Perhaps the most important facet of community ar- chaeology as practiced in Labrador is that it is situated outside the settlements, in the country where the knowl- edge, wisdom, and experience of elders is relevant and ap- parent (Figures 9 and 10). Fieldwork based on mutual re- spect and sharing among families, generations, and visiting researchers honors and encourages indigenous knowledge and different ways of knowing. The practice of commu- nity archaeology with the Innu in Nitassinan is culturally FIGURE 9. No longer the exclusive domain of professional researchers, archaeology in the north has become a cooperative initiative between local community interests and visiting researchers. Here, community activist and former Innu Nation president Daniel Ashini, left, accompanied by Dominique Pokue, survey the ruined shorelines of former Lake Michikamats during an Innu Nation? sponsored archaeological survey of the region in 1995. (S. Loring photograph) 10_Loring_pg115-128_Poles.indd 12410_Loring_pg115-128_Poles.indd 124 11/18/08 9:18:29 AM11/18/08 9:18:29 AM FROM TENT TO TRADING POST AND BACK AGAIN 125 situated experiential education and has an important sub- sistence component. An awareness of animals? especially caribou? trumps the mechanics of fi eldwork: Survey is as much scouting for game as it is searching for the sites where ancestors lived and hunted. The acquisition of game is an integral part of the fi eldwork as young people pre- pare their own trap lines, catch fi sh, and learn from elders how to prepare food and pay proper respect to animals (Loring, 2001b; 2008). And in contrast to the practice of archaeology in the strictly scientifi c paradigm, the les- sons of community archaeology? sharing resources and interpretations and communal decision making? could be neatly summed as ?don?t be bossy, don?t be greedy.? Beyond expanding an awareness and appreciation of in- digenous knowledge and values, Smithsonian archaeology today is about being socially responsible, recognizing that the present is connected to the past, and celebrating indig- enous heritage and land tenure. CONCLUSIONS Around the campfi re or next to the tent stove, the con- versation about archaeology with Native participants is tumbled together with thoughts of the weather, caribou, and seals; of the places where one went hunting, berry picking, or fi shing; and of the old places where ancestors and supernatural creatures once lived. New directions FIGURE 10. Coming full-circle in the production of knowledge about northern people and their history community archaeology returns knowl- edge to a local setting. Working with local youth, and informed by knowledgeable elders, such initiatives serve to celebrate and respect the continuity and experiences of Native northern hosts. Here, visiting elders from Makkovik interpret architectural features at the mid-eighteenth- century Labrador Inuit village site at Adlavik (GgBq-1) in 2002. With them is Lena Onalik (right) from Makkovik, the fi rst professional Inuk archaeologist from Nunatsiavut. (Central Coast of Labrador Archaeological Project photo) 10_Loring_pg115-128_Poles.indd 12510_Loring_pg115-128_Poles.indd 125 11/18/08 9:18:40 AM11/18/08 9:18:40 AM 126 SMITHSONIAN AT THE POLES / LORING in the practice of archaeology in the north recognize the legitimacy of life ?in the country.? Because of different ways of thinking about the past, explaining the past is a basic operating assumption predicated on respect of the cultures and traditions of the people on who live (or used to live) on the land (Lorde, 1981). This collaborative ap- proach of northern anthropological research, predicated on repatriation, recognition, and respect, suggests that the future of the past is likely to re-imagine the cultural and physical landscape of the Arctic in wholly new ways. With the passing of the last vestiges of humanity?s hunt- ing heritage, future generations will need to derive new sources for inspiration. Northern Native involvement with Arctic science might be thought to have begun with the collaboration and insights provided to early explorers and collectors, including those affi liated with the First IPY in 1882? 1883. The increased awareness of the value and acuity of native knowledge and perception has radically transformed the social construction of northern science as the interests and concerns of researchers and indigenous residents alike come to share an interest in the ecological and behavioral consequences of life at high latitudes and a concern for understanding both the past and the future. ACKNOWLEDGMENTS William Fitzhugh (Smithsonian Institution), Igor Krupnik (Smithsonian Institution), and James Fleming (Colby College) provided helpful comments on an earlier draft. LITERATURE CITED Barr, William. 1985. The Expeditions of the First International Polar Year, 1882? 1883. The Arctic Institute of North America, Technical Paper 29. Calgary, Alberta, Canada: University of Calgary. Berger, Thomas R. 1977. Northern Frontier, Northern Homeland: The Report of the Mackenzie Valley Pipeline Inquiry, 2 vols. Toronto: James Lorimer in Association with Publishing Centre, Supply and Services, Canada. 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All early International Polar Year/International Geophysical Year (IPY/ IGY) initiatives were primarily geophysical programs and were exemplary products of the long-established paradigm of ?polar science.? Under that paradigm, scholarly data to be used in academic publications were to be collected by professional scientists and/or by specially trained observers. Arctic indigenous residents had hardly any documented voice in the early IPY/IGY ventures, except by serving as ?subjects? for museum collecting or while working as dog-drivers, guides, and unskilled assistants to research expeditions. Natural scientists with strong interest in Native cultures were the fi rst to break that pat- tern and to seek polar residents as a valuable source of expertise on the Arctic environ- ment. The Smithsonian has a distinguished tradition in working with indigenous experts and documenting their knowledge, from the days of the First IPY 1882? 1883 to the most recent projects on indigenous observations on Arctic climate change. The paper explores the unique role of IPY 2007? 2008 and of recent efforts focused on the documentation of indigenous knowledge of Arctic environment and climate change, by using the experience of one IPY project, SIKU? Sea Ice Knowledge and Use? and research collaboration with local Yupik Eskimo experts from St. Lawrence Island, Alaska. INTRODUCTION This paper explores the emerging links among Arctic people?s ecological knowledge, climate change research, and cultural (or ?social science?) studies in the polar regions. Residents of the Arctic are no strangers to today?s debates about climate change and global warming (Kusugak, 2002; Watt-Cloutier, 2005). Their knowledge on the Arctic environment is being increasingly sought as a source of valuable data for documenting and modeling Arctic climate change (ACIA, 2005). Still, such a rapprochement is not yet an established practice, as many scientists still view Arctic people?s perspectives on climate change as merely ?anecdotal evidence.? Social science?s interest in Arctic people?s observations of climate change is, similarly, a rather recent phenomenon, barely 10 years old (McDonald et al., 1997). Of course, Arctic residents have been observing changes and refl ecting upon fl uctuations in their environment since time immemorial. Their knowl- edge, however, has been ?archived? within northern communities and was transmitted in indigenous languages via elders? stories, personal observations, Igor Krupnik, Department of Anthropology, Na- tional Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 112, Wash- ington, DC 20013-7012, USA (krupniki@si.edu). Accepted 9 May 2008. ?The Way We See It Coming?: Building the Legacy of Indigenous Observations in IPY 2007? 2008 Igor Krupnik 11_Krupnik_pg129-142_Poles.indd 12911_Krupnik_pg129-142_Poles.indd 129 11/17/08 8:34:39 AM11/17/08 8:34:39 AM 130 SMITHSONIAN AT THE POLES / KRUPNIK and information shared among hunters. As scientists be- came increasingly attentive to indigenous perspectives on Arctic climate change, several barriers to productive dialog and communication had to be overcome (see discussion in Krupnik, 2002; Laidler, 2006; Oakes and Riewe, 2006). Over the past decade, this new emerging collaboration produced numerous papers, volumes, collections, docu- mentaries, interactive CD-ROMs, and museum exhibits (Figure 1; Ford et al., 2007; Herlander and Mustonen, 2004; Gearheard et al., 2006; Huntington and Fox, 2005; Krupnik and Jolly, 2002; Laidler, 2006; Laidler and Elee, 2006; Oakes and Riewe, 2006). One of the key tasks of the International Polar Year (IPY) 2007? 2008, articulated in its many documents, is to explore how data generated by polar residents can be matched with the observations and models used by polar scientists (Allison et al., 2007:51? 52; International Coun- cil for Science, 2004:18). For the fi st time, the IPY science program includes a special research theme with a goal to investigate the cultural, historical, and social processes that shape the sustainability of circumpolar human societies, and to identify their unique contributions to global cultural diver- sity and citizenship. (International Council for Science, 2004:15; Krupnik et al., 2005:91? 92) The new IPY includes scores of science projects fo- cused on the documentation of indigenous environmental knowledge and observations of climate change (Hovelsrud and Krupnik, 2006:344? 345; Krupnik, 2007); it serves as an important driver to the growing partnership between Arctic residents and polar researchers. Many Arctic people also see IPY 2007? 2008 as the fi rst international science venture to which they have been invited and one in which FIGURE 1. New public face of polar science, the exhibit Arctic: A Friend Acting Strangely at the National Museum of Natural History, 2006. (Photograph by Chip Clark, NMNH) 11_Krupnik_pg129-142_Poles.indd 13011_Krupnik_pg129-142_Poles.indd 130 11/17/08 8:34:40 AM11/17/08 8:34:40 AM ?THE WAY WE SEE IT COMING?: INDIGENOUS OBSERVATIONS 131 their environmental expertise is valued and promoted. This growing partnership in the documentation of polar residents? observations of climate change is widely viewed as a cutting edge of today?s Arctic social and cultural re- search. SOCIAL SCIENCES IN EARLIER INTERNATIONAL POLAR YEARS All previous International Polar Year initiatives in 1882? 1883, 1932? 1933, and, particularly, the Inter- national Geophysical Year (IGY) in 1957? 1958, were framed primarily, if not exclusively, as geophysical pro- grams focused on meteorology, atmospheric and geomag- netic research, and later, glaciology, geology, space studies, oceanography, and sea ice circulation studies (Fleming and Seitchek, 2009, this volume). We have hardly any record of Arctic residents? involvement in previous IPYs, other than serving as guides, manual laborers, unskilled assistants, or being prospects for ethnographic collecting. None of these earlier IPY ventures organized primarily by meteo- rologists, geophysicists, and oceanographers considered the documentation of indigenous perspectives on Arctic environment a valid topic for scholarly research. Nevertheless, social scientists and polar residents can justly claim a solid IPY legacy of their own that goes back to the fi rst IPY of 1882? 1883. Half of the 12 IPY-1 obser- vational stations and four ?auxiliary? missions that oper- ated in the Arctic produced substantial, often extensive, accounts on local populations and their cultures (Barr, 1985; Krupnik et al., 2005:89? 90). Four seminal ethno- graphic monographs, including three on Arctic indigenous people, were published as direct outcomes of the First IPY (Boas, 1888; Murdoch 1892; Turner 1894), in addition to several chapters in expedition reports, scores of schol- arly articles, and popular accounts (Barr 1985; Burch, 2009, this volume). Some of these contributions? like those by Murdoch and Ray on Barrow; Tromholt (1885) on Kautokeino, Norway; and Bunge (1895) on the Lena River Delta? were illustrated by photographs and draw- ings of local communities, people, and cultural landscapes (Wood and Overland, 2007). Today, such records are trea- sures to museum curators, anthropologists, and histori- ans, but even more so to local communities as resources to their heritage education programs (Crowell, 2009, this volume; Jensen, 2005). Perhaps the most infl uential social science contribu- tion to IPY-1 was the research of Franz Boas, a German- born physicist and, later, the founding fi gure of American anthropology. In 1883, Boas volunteered to do a post-doc study in human geography among the Canadian Inuit as a follow-up to the German IPY-1 observation mission on Baffi n Island (Cole and M?ller-Wille, 1984; M?ller-Wille, 1998). Boas? research among the Baffi n Island Inuit in 1883? 1884 introduced to polar science much of what constitutes today the core of the ?human agenda? of IPY 2007? 2008: the study of indigenous knowledge, adapta- tion, culture change, and Arctic people?s views on the en- vironment. There is no wonder that Boas? monograph on the Central Inuit of Arctic Canada (1888/1964), as well as books by Murdoch on the people of Barrow (1892/1988) and by Turner on the Inuit and Innu of Ungava Bay (1894/2001), remain, perhaps, the most widely cited pub- lications of the entire IPY-1 program. It is also no accident that these IPY-1 monographs on Arctic indigenous people and their cultures were published by the Smithsonian In- stitution. They also remain the only science writings from the First IPY that were ever read and used by Arctic in- digenous people, prior to the recent ?rediscovery? by the Norwegian S?mi of Sophus Tromholt?s photographs of Kautokeino in 1882? 1883. 1 SMITHSONIAN AT THE POLES The Smithsonian Institution has a distinguished record of pioneering cultural research and collecting in the Arc- tic (see Fitzhugh 2002; 2009, this volume). By the time of the fi rst IPY 1882? 1883, the Smithsonian had established productive partnerships with many federal agencies, private parties, and individual explorers (Fitzhugh, 1988b; Loring 2001). The connection to the Signal Offi ce of the War De- partment was essential to the Smithsonian involvement in the fi rst IPY, since the Offi ce was put in charge of the prepa- ration for two U.S. IPY expeditions to Barrow and Lady Franklin Bay in 1881. Dr. Spencer Baird, then Smithsonian Secretary, immediately seized the opportunity to advance the foremost role of the institution in national polar research and to expand its Arctic collections. The Smithsonian was instrumental in selecting natural scientists for both U.S. mis- sions and in training them in conducting observations and collecting specimens. 2 According to the Secretary?s Annual Reports for 1883 and 1884, the Smithsonian assumed re- sponsibility for the natural history component of both U.S. IPY missions and of their collections (Baird 1885a:15? 16; 1885b:15). Baird?s relationship with John Murdoch, one of two natural scientists of the Point Barrow IPY team, is very well documented (Fitzhugh 1988a, xiv? xxix; Murdoch 1892:19? 20). Lucien Turner?s one-man mission to Ungava 11_Krupnik_pg129-142_Poles.indd 13111_Krupnik_pg129-142_Poles.indd 131 11/17/08 8:34:42 AM11/17/08 8:34:42 AM 132 SMITHSONIAN AT THE POLES / KRUPNIK Bay was primarily a Smithsonian (i.e., Baird?s) initiative (Barr, 1985:204; Loring, 2001:xv). To the returning IPY missions, the Smithsonian Insti- tution offered its facilities, libraries, and the expertise of its curators for processing the records and specimens; for these and other efforts the Institution was designated to receive all of the collections brought from the north. The Barrow natural history collections were monumental, as were Turner?s from Labrador. 3 The ethnological portion of the Barrow collection (1,189 specimens upon the origi- nal count 4 ? see Crowell, 2009, this volume) is the sec- ond largest in the National Museum of Natural History (NMNH) Alaska ethnology collections, and Turner?s IPY collection (530 objects) is the second largest among the ethnology acquisitions from Canada. Even most of the ill- fated Greely mission?s natural history specimens, includ- ing some 100 ethnological objects (Greely, 1888:301? 317) and personal memorabilia, ended up in the Smithsonian collections (Neighbors, 2005). All three U.S. IPY missions also produced several dozen photographs that are among the earliest from their respective areas. 5 By the very scope of their assignments, early IPY sci- entists combined instrumental meteorological observa- tions with natural history research and collecting; hence, the changes in local climate and natural environment may have been on their minds as well. We know that Boas was deeply interested in Inuit perspectives on their envi- ronment. He had been systematically documenting Inuit knowledge of sea ice, snow, weather, place-names, and navigation across the snow/ice covered terrain as part of his research program (Cole and M?ller-Wille, 1984:51? 53), very much like many IPY scientists are doing today. In 1912, 30 years after Murdoch and Turner, Smith- sonian anthropologist Riley Moore visited St. Lawrence Island, Alaska, and worked with a young hunter named Paul Silook (Figure 2). Silook assisted Moore in translating elders? stories about the famine of 1878? 1879 that killed hundreds of island residents (Moore, 1923:356? 358). Some scientists believe that the famine was caused by ex- traordinary sea ice and weather conditions that disrupted the islanders? hunting cycle (Crowell and Oozevaseuk, 2006). In the late 1920s, another Smithsonian scientist, Henry Collins, partnered with Silook in search for local knowledge about the early history of island?s population. Collins also asked Silook to maintain a personal diary with the records of weather conditions; this diary has been pre- served at the Smithsonian National Anthropological Ar- chives (Jolles, 1995). Silook may have told Collins about the catastrophic storms that destroyed his native village of Gambell in 1913 and other extraordinary events, which he described to other scholars in later years (Krupnik et al., 2002:161? 163). CONVERTING LOCAL OBSERVATIONS INTO ?IPY SCIENCE? If partnership between Arctic residents and polar re- searchers in IPY 2007? 2008 is to bring tangible benefi ts to both sides, each party has to understand how the other observational system works. This implies certain steps needed to make the two systems compatible or, at least, open to data exchange. Local knowledge, very much like science, is based upon long-term observation and moni- toring of dozens of environmental parameters, in other FIGURE 2. Paul Silook, Siluk (1892? 1946), worked with many sci- entists who came to his home village of Gambell over more than three decades, between 1912 and 1949. (Photograph by Riley D. Moore, 1912, Smithsonian Institution. NAA, Neg. # SI 2000-693) 11_Krupnik_pg129-142_Poles.indd 13211_Krupnik_pg129-142_Poles.indd 132 11/17/08 8:34:43 AM11/17/08 8:34:43 AM ?THE WAY WE SEE IT COMING?: INDIGENOUS OBSERVATIONS 133 words, upon multifaceted data collection. By and large, indigenous experts follow many of the same analytical steps, though in their specifi c ways (Berkes 1999:9? 12). Much like scientists, local hunters exchange individual observations and convert them into a shared body of data. They analyze the signals of change and seek expla- nations to the phenomena they observe (Krupnik 2002; Huntington et al., 2004). When the fi rst projects in the documentation of indig- enous observations of Arctic climate change were started, scientists were literally overwhelmed by the sheer wealth of local records. As a result, much of the early work on indigenous observations, up to 2003? 2005, focused on the mere documentation of various evidence of change coming from different areas. 6 Next, scientists tried to apply cer- tain tools, such as typologies, maps, and matrix tables ar- ranged by ecosystem component, to compare reports from different areas (McDonald et al., 1997:46? 47; Krupnik and Jolly, 2002; Huntington and Fox, 2005). These fi rst applications of scientifi c tools illustrated that Arctic resi- dents observe a consistent pattern of change and that they interpret the phenomena they observe in a comprehensive, integrated manner. It also became clear that local people have documented rapid change in the Arctic environment in a profound and unequivocal way. The next step in scientists? approach to indigenous re- cords is to look for cases and areas where indigenous and scientifi c data disagree and offer differing, often confl ict- ing interpretations (Huntington et al., 2004; Krupnik and Ray, 2007; Norton, 2002). This approach reveals certain features of indigenous versus scientifi c observation pro- cesses, such as differences in scaling, in the use of prime in- dicators, and in causes and linkages cited as explanations in two knowledge systems. It also offers a much more sys- temic vision that goes beyond a popular dichotomy that contrasts local or traditional ecological knowledge (TEK) and the scientifi c knowledge. Under such vision, the for- mer is usually labeled ?intuitive, holistic, consensual, and qualitative,? whereas the latter is perceived as analytical, quantitative, and compartmentalized (Bielawski, 1992; Krupnik, 2002:184). While these labels contain some truth, Native experts have demonstrated repeatedly that they can effectively operate with both types of records and that they often match them more skillfully than scien- tists do (Aporta and Higgs, 2005; Bogoslovskaya, 2003; Krupnik and Ray, 2007; Noongwook et al., 2007). Scientists commonly argue that Arctic people?s records of climate change would be a valuable contribution to IPY 2007? 2008 (Allison et al., 2007). Still, such accommoda- tion requires substantial mutual adjustment of observa- tional and analytical practices. Scientists have to accept that data generated by local observers are crucial to cover certain gaps in instrumental or satellite records, despite some reservations with regard to how local observations are collected and transmitted. From their side, Native ex- perts participating in joint projects have to acknowledge certain standards of science data collection, like consis- tency, transparency, and independent verifi cation. Here the gap is indeed serious, since indigenous observations are mostly non-numerical, are freely and widely shared within the community, and are rarely if ever reported in writing (Bates, 2007:89? 91). Because of these and other factors, indigenous records can rarely be tested by scien- tists? analytical procedures, like long-term series, statisti- cal averaging, correlation, and trend verifi cation, among others. Also, indigenous observers have their specifi c ?terms of references? when assessing the validity of their data, such as individual life experience, community-based mem- ory, or verifi cation by elders or individual experts (Noong- wook et al., 2007:48; Gearheard et al., 2006). Neverthe- less, the sheer volume of data to be generated by many participatory projects in IPY 2007? 2008 has already trig- gered efforts to develop procedures and standards for local observations and for management of indigenous records. 7 SIKU? SEA ICE KNOWLEDGE AND USE The experience of one such project illustrates what scientists can learn from local experts and how indigenous knowledge may advance IPY science. ?Sea Ice Knowl- edge and Use: Assessing Arctic Environmental and Social Change? (SIKU, IPY #166) is an IPY project aimed at the documentation of indigenous observations of Arctic cli- mate change, with its focus on sea ice and the use of ice- covered habitats by polar residents. The project?s acronym SIKU is also the most common word for sea ice (siku) in all Eskimo languages, from Bering Strait to Greenland. As a collaborative initiative, SIKU relies on partnership among anthropologists, geographers, and marine and ice scientists from the United States, Canada, Russia, Green- land, and France, and indigenous communities in Alaska, Canada, Greenland, and Russian Chukotka. SIKU is orga- nized as a consortium of several research initiatives sup- ported by funds from various national agencies. The proj- ect was started in winter 2006? 2007 and it will continue through 2008 and 2009. The Alaska-Chukotka portion of SIKU has its three hubs at the Smithsonian Arctic Studies Center (managed by Igor Krupnik), the Russian Institute 11_Krupnik_pg129-142_Poles.indd 13311_Krupnik_pg129-142_Poles.indd 133 11/17/08 8:34:45 AM11/17/08 8:34:45 AM 134 SMITHSONIAN AT THE POLES / KRUPNIK of Cultural and Natural Heritage in Moscow (Lyudmila Bogoslovskaya), and the University of Alaska, Fairbanks (Hajo Eicken). The Canadian portion of SIKU is called Inuit Sea Ice Use and Occupancy Project (ISIUOP); it is co- ordinated by Claudio Aporta and Gita Laidler at Carleton University, Ottawa (see http://gcrc.carleton.ca/isiuop). Research under the SIKU-Alaska and SIKU- Chukotka program takes place in several local communities, such as Gambell, Shaktoolik, Wales, Shishmaref, Barrow, Tununak, Uelen, Lavrentiya, Sireniki, and so on (Figure 3). It includes daily ice and weather observations, collections of Native terms for sea ice and weather phenomena, docu- mentation of ecological knowledge related to sea ice and ice use from elders and hunters, and searches for historical records of ice and climate conditions (see http://www.ipy .org/index.php?ipy/detail/sea_ice_knowledge_and_use/). This paper examines the contribution of one of such lo- cal SIKU observers, Leonard Apangalook Sr., a hunter and community leader from the Yupik village of Gambell on St. Lawrence Island, Alaska. Apangalook, 69, is a nephew of Paul Silook and he continues an almost 100-year tra- dition of his family?s collaboration with Smithsonian scientists (Figure 4). Since spring 2006, Apangalook has produced daily logs on sea ice, weather, and local subsis- tence activities in his native community of Gambell. His personal contribution to the IPY 2007? 2008 now covers two full ?ice years,? 2006? 2007 and 2007? 2008, and will hopefully extend into 2008? 2009. St. Lawrence Island residents? knowledge of sea ice has been extensively documented in recent years via sev- eral collaborative projects with two local Yupik communi- ties of Gambell and Savoonga (Huntington, 2000; Jolles, FIGURE 3. The Yupik village of Gambell on St. Lawrence Island, Alaska, is one of the key research ?sites? for the SIKU project. (Photo, Igor Krupnik, February 2008) 11_Krupnik_pg129-142_Poles.indd 13411_Krupnik_pg129-142_Poles.indd 134 11/17/08 8:34:45 AM11/17/08 8:34:45 AM ?THE WAY WE SEE IT COMING?: INDIGENOUS OBSERVATIONS 135 1995; 2003; Krupnik, 2000; 2002; Krupnik and Ray, 2007; Metcalf and Krupnik, 2003; Noongwook et al., 2007; Oozeva et al., 2004). The island people have long voiced concerns about shifts in the local environment they observed and about the growing impact of climate change upon their economy and way of living. Apangalook?s ob- servations help convert these statements into a written re- cord open to scientifi c scrutiny and analysis. NEW PATTERNS OF FALL ICE FORMATION St. Lawrence Islanders have a highly nuanced vision on how the new sea ice is being formed in their area or, rather, how it used to be formed in the old days. Their native Yupik language has more than 20 terms for various types of young ice and freezing conditions. Winds and currents would drive small chunks of fl oating ice (kulusiit) from the north in October or even in late September (Oozeva et al., 2004:133? 134). Most would melt or would be washed ashore; but some would freeze into the locally formed slush or frazil ice. Around the time the fi rst local ice is established (in late October or early November), the prevailing winds would shift direction, from primarily southerly to north- erly, followed by a drop in daily temperatures. Depending upon year-to-year variability, but usually by late November, the thick Arctic ice pack, sikupik, would arrive from the Chukchi Sea, often smashing the young locally formed ice. Crashing, breaking, and refreezing would continue through December, until a more solid winter ice was formed to last until spring (Oozeva et al., 2004:136? 137). Since the 1980s, hunters started to observe changes to this pattern, which had until then been seen as normal. First, drifting ice fl oes, kulusiit, were late to arrive, often by a full month, and by the late 1990s, they have stopped coming altogether. The new ice is now being formed en- tirely out of local frozen slush or frazil ice, via its thick- ening, consolidation, repeated break-ups, and refreezing. Then the main pack ice ceased to arrive until January or even February; and in the last few years it did not arrive at all. Even when the pack ice fi nally comes from the north, it is not a solid thick ice, sikupik, but rather thin new ice that was formed further to the north. Because of this new set of dynamics, the onset of winter ice conditions on St. Lawrence Island is now delayed by six to eight weeks, that is, until late December or even January. 8 Apangalook?s observations during two IPY winters of 2006? 2007 and 2007? 2008 help document this new pattern in great detail. In addition, his daily records may be matched with the logbooks of early teachers from his village of Gambell that covered three subsequent win- ters of 1898? 1899, 1899? 1900, and 1900? 1901 (Doty, 1900:224? 256; Lerrigo, 1901:114? 132; 1902:97? 123; see Oozeva et al., 2004:168? 191). According to Apangalook?s logs, no drifting ice fl oes were seen in winter 2006? 2007 and none in the month of November 2007. Although in 2007? 2008 the formation of slush ice started more than two weeks earlier than in 2006? 2007, the ice was quickly broken up by a warming spell, so that on 22 November 2007, Apangalook reported: ?Thanksgiving Day with no ice in the ocean; normally [we] would have ice and hunt walrus on Thanksgiving Day but not anymore.? In both 2006 and 2007, the temperature dropped solidly below freezing on the fi rst week of December? and that was two to four weeks later than a century ago. The change of wind regime, from southerly to predominately northerly winds, also occurred in early December, that is, two to four weeks later than in 1898? 1901 (Oozeva et al., 2004:185). Due to shift in wind and temperature, local slush ice solidifi ed rapidly. On 16 December 2006, Apangalook reported that ?when locally formed ice gets thick and encompasses the entire Bering Sea around our island, our elders used to say that we have a winter that is locally formed? (Figure 5). WINTER WEATHER AND ICE REGIMES Apangalook?s daily logs substantiate statements of other St. Lawrence hunters about the profound change in winter weather and ice regime over the past decades (cf. Oozeva et al., 2004; Noongwook et al., 2007). With the FIGURE 4. Leonard Apangalook Sr., SIKU participant and local observer in the village of Gambell, St. Lawrence Island (courtesy of Leonard Apangalook). 11_Krupnik_pg129-142_Poles.indd 13511_Krupnik_pg129-142_Poles.indd 135 11/17/08 8:34:56 AM11/17/08 8:34:56 AM 136 SMITHSONIAN AT THE POLES / KRUPNIK absence of solid pack ice, people have to adapt to a far less stable local new ice that can be easily broken by heavy winds, storms, and even strong currents (Figure 6). Accord- ing to the elders, this ice is ?no good,? as it is very dangerous for walking and winter hunting on foot or with skin boats being dragged over it, as used to be a common practice in the ?old days? (Oozeva et al., 2004:137? 138, 142? 143, 163? 166). As a result, few hunters dare to go hunting on ice in front of the village in wintertime (Figure 7). Instead, they have to rely upon shooting the animals from the shore or from ice pressure ridges (Figure 8) or upon hunting in boats in the dense fl oating ice, a technique that is now the norm in Gambell during winter months (Figure 9). In the early teachers? era of 100 years ago, boat hunting for wal- ruses did not start in Gambell until early or even late March (Oozeva et al., 2004:187? 188). Winter conditions in Gambell, with the prevailing northerly winds, are often interrupted by a few days of vi- FIGURE 5. The new ice is being formed from the pieces of fl oat- ing icebergs and newly formed young ice. There are terms for every single piece of ice in this picture and many more in the local language (?Watching Ice and Weather Our Way,? 2004, p. 114; original photo by Chester Noongwook, 2000). FIGURE 6. ?Winter that is locally formed.? Thin young ice solidi- fi es along the shores of St. Lawrence Island, February 2008. Leon- ard Apangalook stands to the right, next to his sled. (Photo, Igor Krupnik, 2008) FIGURE 7. Hunters rarely venture onto young unstable ice and they usually prefer to be accompanied by an experienced senior person. (Photo, Hiroko Ikuta, 2006) FIGURE 8. Two Gambell hunters are looking for seals from ice pres- sure ridges. The ridges look high and solid, but they can be quickly destroyed under the impact of strong wind or storms. (Photo, Igor Krupnik, February 2008) 11_Krupnik_pg129-142_Poles.indd 13611_Krupnik_pg129-142_Poles.indd 136 11/17/08 8:34:59 AM11/17/08 8:34:59 AM ?THE WAY WE SEE IT COMING?: INDIGENOUS OBSERVATIONS 137 olent storms and warm spells brought by southerly winds. This happened twice in the winters of 1899? 1900 and 1900? 1901, and three times in the winter of 1898? 1899 (Oozeva et al., 2004:185). According to Conrad Oozeva, an elderly hunter from Gambell, the warm spells have been also typical in his early days: We commonly have three waves of warm weather and thaw- ing during the wintertime. After these warmings, we love to go hunting in boats on water opening, before it covers again with the new ice. The only difference I see is that these warm waves were not long enough, just a few days only. We now have lon- ger warming waves during the winter, often for several days. (Oozeva et al., 2004:186) These days, violent winter storms often lead to nu- merous episodes of ice breakups and new ice formation every winter. Apangalook?s record indicates at least four episodes of complete ice disintegration in Gambell in the period from December 2006 until March 2007. On 10 January 2007, he wrote in his log, ?It is unusual to have swells and to lose ice in January compared to normal years in the past. Locally formed ice that covered our area to 9/10 easily disappears with rising temperatures and storm generated swells.? Then on 31 January 2007, he reported again: What a twist we have in our weather situation at the end of January! Wind driven waves cleared away pressure ridges on west side with open water west of the Island. Part of the shore- fast ice broke away on the north side beach also from the swells. Unusual to have so many low pressures channel up the Bering Straits from the south in a sequence that brought in high winds and rain. Much of the snow melted, especially along the top of the beach where we now have bare gravel. Undeniably, the cli- mate change has accelerated over the past fi ve years where se- verity of winds and erratic temperatures occur more frequently every year. Elders on St. Lawrence Island and in other north- ern communities unanimously point toward another big change in winter conditions. In the ?old days,? despite episodic snowstorms and warm spells, there were always extended periods of quiet, cold weather, with no winds. These long cold stretches were good for hunting and trav- eling; they also allowed hunters to predict the weather and ice conditions in advance. In the teachers? records of more than a century ago, those stretches of calm cold weather often covered two to three weeks. This pattern does not occur today. According to Apangalook?s logs, there were a few periods of relatively quiet weather during the winter of 2006? 2007; but they lasted for a few days only. Decem- ber 2006 was particularly unstable and windy, with just one calm day. In comparison, during December 1899, the weather was quiet for 18 days, in two long stretches. In December 1900, quiet and calm weather persisted for al- most 10 days in a row (Oozeva et al., 2004:185? 186). No wonder Arctic elders claim that ?the earth is faster now? (cf. Krupnik and Jolly, 2002:7). CHANGES IN MARINE MAMMAL BEHAVIOR AND HABITATS Local hunters? observations are naturally fi lled with the references to wildlife and subsistence activities. When seen upon a broader historical timeframe, those records provide compelling evidence of a dramatic shift that is tak- ing place in the northern marine ecosystems. Apangalook reported (6 March 2007): A few years back when the polar pack ice did not reach our area anymore, we sighted bowhead whales in our area spo- radically in the middle of winter. Back when our seasons were normal, we saw whales in the fall going south for the winter and didn?t see any in mid-winter, until they start coming back in mid-March-April and May. Now, with more whales in our area in mid-winter we know that they are (mostly) wintering in our area. Without polar pack ice we had suspected that some of the whales stopped that migration north of our island and are not going further south anymore. . . . We know today that their wintering area is further north. FIGURE 9. Hunting in boats in dense fl oating ice is now a com- mon practice in Gambell during the wintertime. (Photo, G. Carleton Ray) 11_Krupnik_pg129-142_Poles.indd 13711_Krupnik_pg129-142_Poles.indd 137 11/17/08 8:35:08 AM11/17/08 8:35:08 AM 138 SMITHSONIAN AT THE POLES / KRUPNIK Many local hunters, like Apangalook, argue that whale migrations have been deeply affected by climate change. Hunters started observing bowhead whales off St. Law- rence Island in December (usually, a few animals) since 1962. Since 1995, winter whaling has become a common practice in both Gambell and the other island community of Savoonga, so that some 40 percent of bowhead whales are now being taken in wintertime (Noongwook et al., 2007:51). The difference is indeed remarkable compared to the conditions of a century ago, when the bowhead whales were not hunted in Gambell until early April (as in 1899) or even early May, as in 1900 and 1901 (Oozeva et al., 2004:189). To the contrary, in early 2007, Apangalook re- ported sightings of bowhead whales on February 6, and the hunters fi rst tried to pursue them on 10 February 2007. The whales were then seen off Gambell repeatedly for the entire month. Local hunters in Barrow have also spotted bowhead whales in mid-February 2007; recent underwa- ter acoustic recording off Barrow documented the presence of gray whales (Eschrichtius robustus) during the winter of 2003/2004 (Stafford et al., 2007:170). If whales have become frequent in the northern Bering Sea and southern Chukchi Sea in mid-winter, that would be a strong indica- tor to the dramatic shift in their migration and distribution pattern across the North Pacifi c? Western Arctic region. Pacifi c walrus, another marine mammal species of critical importance to St. Lawrence Island hunters is also becoming more common in wintertime. Back in the ?olden days,? walrus used to come to the island in great numbers in late October or early November, ahead of the moving polar pack ice (Krupnik and Ray, 2007:2950). The bulk of the herd usually moved south of the island in December; but a small number of bull walruses commonly remained around Gambell in wintertime, mostly in offshore leads and polynyas. Still, the main hunting season for walrus did not start until late April or May. According to Apan- galook?s records, the fall arrival of walrus in both 2006 and 2007 was delayed by several weeks, so that the fi rst walruses were not seen until mid-December. After that, walruses have been hunted in Gambell on almost daily ba- sis throughout the winter of 2006? 2007, which indicates that, like bowhead whales, they are also staying in grow- ing numbers during the winter months. Besides ?wintering? walruses, Gambell hunters in win- ter 2007? 2008 have observed several ribbon seals (Phoca fasciata) that are normally not seen in the area until late April or May. In February 2007, SIKU observers in Wales, some 200 miles to the north of Gambell, have spotted belugas (white whales); in the ?old days,? beluga whales had not been seen in the Bering Strait area until mid-April. Whereas some species are becoming more common in winter, others are moving out. Apangalook reports: One species of birds we used to hunt ever since I was a young child was the old squaw [Clangula hyemalis? IK]. Flocks of these birds in the thousands would fl y north every morning at daybreak and return in the evening to [the] leeward side of the island. This pattern of movement daily I have seen every day in the sixty plus years of my life; but within the past fi ve years the numbers have dwindled to near zero. [. . .] Are their food sources moving to a different area or is [it] getting depleted? (1 February 2007) CONCLUSIONS: MESSAGES FROM INDIGENOUS OBSERVATION The evidence from the fi rst systematic monitoring by lo- cal IPY observers confi rms a substantial shift in sea ice and weather regime over that past decade, as has been claimed by polar scientists and indigenous experts alike. It is char- acterized by a much shorter presence of sea ice, often by several weeks. Many types of ice are becoming rare or have completely disappeared from the area, such as solid pack ice or forms of multi-layered ice built of old and new ice. Most of the ice formations are now built of young and frag- ile fi rst-year ice. The ice is also becoming increasingly un- stable and dangerous for hunters. 9 In Apangalook?s words, ?Even marine mammals avoid this kind of ice condition, as it is hazardous to the animals too. It looks like game animal are taking refuge in more solid ice elsewhere.? Hunters? observations also confi rm the profound northward shift in the Bering Sea marine ecosystem, which is also detected by scientists (Grebmeier et al., 2006; Lit- zow, 2007). Arctic residents refer to such shift in many of their descriptions of climate change, although they use their own ?fl agship species? as prime indicators. In- digenous hunters, naturally, pay most attention to large marine mammal and bird species as opposed to fi sh, inver- tebrates, and benthic communities that are popular indica- tors of change among marine biologists and oceanogra- phers (Grebmeier et al., 2006; Ray et al., 2006; Sarmiento et al., 2004). Arctic residents are extremely worried about the im- pact of ecosystem change on their economies, culture, and lifestyles. As Apangalook put it, Predictability of our game animals of the sea are inconsis- tent and erratic compared (to) how it used to be back in the nor- mal seasons. We, the hunters, along with the marine mammals 11_Krupnik_pg129-142_Poles.indd 13811_Krupnik_pg129-142_Poles.indd 138 11/17/08 8:35:11 AM11/17/08 8:35:11 AM ?THE WAY WE SEE IT COMING?: INDIGENOUS OBSERVATIONS 139 we hunt, are truly at the mercy of our rapidly changing environ- ment. (Apangalook, 2 March 2007) The last message from indigenous records is that local observations could be a valuable component of any instru- mental observation network built for IPY 2007? 2008 and beyond. Many hunters in small Alaskan villages are trained to keep daily weather logs and are very familiar with the practices of instrumental observation and forecasting. Also, their daily records can be matched with historical instru- mental data from the same areas that sometimes go back to the years of the First IPY of 1882? 1883 (Wood and Over- land 2006), as well as with the readings of today?s weather stations and ice satellite imagery. Such cross-reference with the long-term instrumental series would create the needed comparative context to indigenous observations and would help introduce analytical scholarly tools to the analysis of indigenous data. Arctic residents? observations are too pre- cious a record to be discounted as ?anecdotal evidence? in today?s search for the documentation and explanation of environmental change. Besides, local observations in places like Gambell, with the now-shortened ice season and thinned fi rst-year ice, may offer a valuable insight into the future status of the Arctic sea ice of many climate models. Those mod- els predict the shrinking, thinning, and eventual loss of multi-year ice over almost the entire Arctic Ocean by the middle of this century (Bancroft, 2007; Johannessen et al., 2004:336? 338; Overland, 2007; Richter-Menge et al., 2006). Arctic residents? integrative vision of their environ- ment can be invaluable to our understanding of this new Arctic system in the decades to come. ACKNOWLEDGMENTS This paper is a tribute to the long-term partnership with local experts in the documentation of their observa- tions of sea ice and climate change in the Bering Strait region. Collaboration with Herbert Anungazuk, Leonard Apangalook Sr., George Noongwook, Chester Noong- wook, Conrad Oozeva, Willis Walunga, Winton Weyapuk Jr., and others turned into the bonds of friendship and gained new momentum under the SIKU project in 2006? 2008. I am grateful to Leonard Apangalook Sr.; also to Hiroko Ikuta, Chester Noongwook, and G. Carleton Ray, who shared their photos for illustrations to this paper. My colleagues Ernest S. Burch Jr., Aron Crowell, William Fitzhugh, Molly Lee, G. Carleton Ray, Cara Seitchek, Dennis Stanford, and Kevin Wood offered helpful com- ments to the fi rst drafts of this paper. I thank them all. NOTES 1. Sophus Tromholt?s photographs taken in 1882? 1883 have been displayed at the ?Indigenous Opening? of IPY 2007? 2008 and used in a special trilingual calendar produced for the event. See http://www.ip-py.org/ news_cms/2007/january/tromholdt_exhibit_at_the_opening_ ceremony/6 (accessed 30 March 2008). 2. Point no. 51 in Hazen?s instructions, ?Observations and col- lections in the realms of zoology, botany, geology, &c.? (Ray, 1885:13; Greely, 1888:104). 3. The Barrow collections included ?497 bird-skins, comprising about 50 species, and 177 sets of eggs; [. . .] a small collection of skins, skulls, and skeletons of mammals; 11 or 12 species of fi shes; a very few insects; and some marine and fresh-water invertebrates. The plants of the region were carefully collected. A considerable number of Eskimo vo- cabularies were obtained, together with a large collection of implements, clothing, &c? (Baird, 1885a:15). Turner?s collections from Labrador were described as ?[. . .] of birds, 1,800 specimens; eggs, 1,800 speci- mens; fi shes, 1,000 specimens; mammals, 200 specimens; ethnological, 600 artifacts; plants, a great number; insects, over 200,000; geological specimens, a great variety; Eskimo linguistics, over 500 pages of manu- script, embracing thousands of words and over 800 sentences? (Baird, 1885b:17). 4. In fact, there are 1,068 ethnological objects, according to today?s electronic catalog, with Lt. P. Ray recorded as donor; plus 6 objects do- nated by Capt. Herendeen, another member of the Barrow mission. 5. Barrow team also conducted a census of the residents of Bar- row, with 137 names of men, women, and children (Ray, 1885:49); that makes it one of the earliest samples of personal Inuit names from Arctic Alaska. 6. See reviews of several individual documentation projects on in- digenous observations in Krupnik and Jolly, 2002; also Herlander and Mustonen, 2004; Huntington and Fox, 2005. 7. One of such projects, ELOKA (Exchange for Local Observa- tions and Knowledge of the Arctic, IPY # 187) works ?to provide data management tools and appropriate means of recording, preserving, and sharing data and information? from Arctic communities? See http:// nsidc.org/eloka/ (accessed 30 March 2008). 8. This pattern has been consistently reported by indigenous ob- servers across the Arctic area? see Gearheard et al., 2006; Laidler, 2006; Laidler and Elee, 2006; Laidler and Ikummaq, 2008; McDonald et al., 1997; Metcalf and Krupnik, 2003. 9. See similar conclusions from other projects in the documenta- tion of indigenous knowledge on sea ice change (Gearheard et al., 2006; Laidler, 2006; Laidler and Elee, 2006; Norton, 2002). Murdoch?s work in Barrow even inspired a special IPY 2007? 2008 project aimed at repli- cating his ethnological collections by today?s specimens (Jensen, 2005) LITERATURE CITED ACIA. 2005. Arctic Climate Impact Assessment (ACIA). Cambridge, U.K.: Cambridge University Press. Allison, I., M. B?land, K. Alverson, R. Bell, D. Carlson, K. Darnell, C. Ellis-Evans, E. Fahrbach, E. Fanta, Y. Fujii, G. Glasser, L. Goldfarb, G. Hovelsrud, J. Huber, V. Kotlyakov, I. Krupnik, J. Lopez- Martinez, T. Mohr, D. Qin, V. Rachold, C. Rapley, O. Rogne, E. Sarukhanian, C. Summerhayes, and C. Xiao. 2007. The Scope of Science for the International Polar Year 2007? 2008. World Meteorological Organi- zation, Technical Documents 1364. Geneva. Aporta, C., and E. Higgs. 2005. Satellite Culture: Global Positioning Systems, Inuit Wayfi nding, and the Need for a New Account of Technology. 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E. Overland. 2006. Climate Lessons from the First International Polar Year. Bulletin of the American Meteorological Society, 87(12): 1685? 1697. ???. 2007. Documentary Image Collection from the First Interna- tional Polar Year, 1881? 1884. http://www .arctic .noaa .gov/ aro/ ipy- 1/ Frontpage.htm (accessed 3 September 2007). 11_Krupnik_pg129-142_Poles.indd 14211_Krupnik_pg129-142_Poles.indd 142 11/17/08 8:35:13 AM11/17/08 8:35:13 AM ABSTRACT. In the Southern Ocean, 205 species of pelagic calanoid copepods have been reported from 57 genera and 21 families. Eight species are found in the coastal zone; 13 are epipelagic, and 184 are restricted to deepwater. All 8 coastal species and eight of 13 epipelagic species are endemic, with epipelagic species restricted to one water mass. Of the 184 deepwater species, 50 are endemic, and 24 occur south of the Antarctic Convergence. Most of the remaining 134 deepwater species are found throughout the oceans with 86% percent reported as far as the north temperate region. The deepwater genus Paraeuchaeta has the largest number of species in the Southern Ocean, 21; all are carnivores. Scolecithricella is also speciose with 16 species, and more specimens of these detritivores were collected. Species with a bipolar distribution are not as common as bipolar species pairs. A bipolar distribution may result from con- tinuous extinction in middle and low latitudes of a wide spread deepwater species with shallow polar populations. Subsequent morphological divergence results in a bipolar species pair. Most of the numerically abundant calanoids in the Southern Ocean are endemics. Their closest relative usually is a rare species found in oligotrophic habitats throughout the oceans. Abundant endemics appear adapted to high primary and sec- ondary productivity of the Southern Ocean. Pelagic endemicity may have resulted from splitting a widespread, oligotrophic species into a Southern Ocean population adapted to productive habitats, and a population, associated with low productivity that remains rare. The families Euchaetidae and Heterorhabdidae have a greater number of their endemic species in the Southern Ocean. A phylogeny of these families suggests that independent colonization by species from different genera was common. Thus, two building blocks for the evolution of the Southern Ocean pelagic fauna are independent colonization and adaptation to high productivity. INTRODUCTION Copepods often are referred to as the insects of the seas. They certainly are comparable to insects in survival through deep time, ecological dominance, geo- graphic range, and breadth of adaptive radiation (Schminke, 2007). However, they are not comparable to insects in numbers of species. Only 11,302 species of copepods were known to science toward the end of the last century (Humes, 1994), and 1,559 have been added since then. In contrast, the number of de- scribed insects approaches one million (Grimaldi and Engel, 2005). In terms of the number of individuals alive at any one time, however, copepods undoubtedly E. Taisoo Park, Texas A&M University, 29421 Vista Valley Drive, Vista, CA 92084, USA. Frank D. Ferrari, Invertebrate Zoology, Na- tional Museum of Natural History, Smithsonian Institution, 4210 Silver Hill Road, Suitland, MD 20746, USA. Corresponding author: F. D. Ferrari ( ferrarif@si.edu). Accepted 27 June 2008. Species Diversity and Distributions of Pelagic Calanoid Copepods from the Southern Ocean E. Taisoo Park and Frank D. Ferrari 12_Park_pg143-180_Poles.indd 14312_Park_pg143-180_Poles.indd 143 11/17/08 8:30:12 AM11/17/08 8:30:12 AM 144 SMITHSONIAN AT THE POLES / PARK AND FERRARI surpass the insects. Among the copepod orders, calanoid copepods contribute more numbers of individuals to the Earth?s biomass, primarily because of their unique success in exploiting pelagic aquatic habitats. Calanoid copepods also are speciose; Bowman and Abele (1982) estimated 2,300 species of calanoids, and as of this writing, 525 spe- cies have been added. These calanoid species are placed in 313 nominal genera belonging to 45 families (F. D. Ferrari, personal database). Knowledge about the distribution and diversity of calanoid copepods in the Southern Ocean has increased signifi cantly over the past century (Razouls et al., 2000). Most of the calanoid copepods reported from the South- ern Ocean have been collected from pelagic waters. How- ever, more species new to science are now being described from waters immediately over the deep-sea fl oor of the Southern Ocean (Bradford and Wells, 1983; Hulsemann, 1985b; Schulz and Markhaseva, 2000; Schulz, 2004, 2006; Markhaseva and Schulz, 2006a; Markhaseva and Schulz, 2006b, 2007a, 2007b). The diversity of this ben- thopelagic calanoid fauna from other oceans (Grice, 1973; Markhaseva and Ferrari, 2006) suggests that the total cal- anoid diversity from this habitat of the Southern Ocean is signifi cantly underestimated, and many new species are expected to be described. The present review then is restricted to pelagic calanoid copepods because the ben- thopelagic fauna has not been well surveyed and their spe- cies not as well known as pelagic calanoids. Pelagic calanoid copepods are numerically the domi- nant species of the zooplankton community in the South- ern Ocean (Foxton, 1956; Longhurst, 1985). Beginning with the Challenger expedition (1873? 1876), many expe- ditions to the Southern Ocean have provided specimens for taxonomic studies of the calanoids. Early works by Brady (1883), based on the Challenger collections, Gies- brecht (1902), based on Belgica collections, Wolfenden (1905, 1906, 1911), based on the Gauss (German deep- sea expedition) collections, and Farran (1929), based on the British Terra Nova collections, led to the discovery of most of the numerically dominant and widespread pelagic calanoid species in the Southern Ocean. Several major national expeditions followed, such as the Me- teor expedition, 1925? 1927, the SS Vikingen expedition, 1929? 1930, and the Norvegia expedition, 1930? 1931. However, collections obtained by these expeditions were studied mainly to understand the vertical or seasonal distribution or other aspects of the biology of pelagic animals. Signifi cant publications resulting from these studies include Hentschel (1936), Steuer and Hentschel (1937), and Ottestad (1932, 1936). In 1925 the British Discovery Committee launched a program of extensive oceanographic research in the Southern Ocean, includ- ing intensive studies of the zooplankton fauna. Publica- tions by Mackintosh (1934, 1937), Hardy and Gunther (1935), and Ommanney (1936) based on the Discovery collections are notable for their valuable contributions to the population biology of the numerically dominant cala- noid copepods. Continued studies of the Discovery col- lections led to the publication of additional papers, such as Foxton (1956) about the zooplankton community and Andrew (1966) on the biology of Calanoides acutus, the dominant herbivore of the Southern Ocean. Southern Ocean copepods became the subject of taxo- nomic studies once again toward the middle of the last century with two important monographs (Vervoort, 1951, 1957). These were the most comprehensive treatments published on pelagic calanoids, to that time, and began a new era of taxonomic analyses of Southern Ocean cope- pods. In these two studies, many previously known species of the Southern Ocean calanoids were completely and care- fully redescribed, confusion regarding their identity was clarifi ed, and occurrences of these species in other oceans were noted from the published literature. Two papers by Tanaka (1960, 1964) appeared soon afterward, reporting on the copepods collected by the Japanese Antarctic Expe- dition in 1957 and 1959. On the basis of collections made by the Soviet Antarctic expeditions, 1955? 1958, Brodsky (1958, 1962, 1964, 1967) published several studies of the important herbivorous genus Calanus. More recently, im- portant contributions to taxonomy of the Southern Ocean calanoids have been made by Bradford (1971, 1981) and Bradford and Wells (1983), reporting on calanoids found in the Ross Sea. Additionally, invaluable contributions have been made to the taxonomy of the important in- shore genus Drepanopus by Bayly (1982) and Hulsemann (1985a, 1991). Beginning in 1962, the U.S. Antarctic Research Pro- gram funded many oceanographic cruises to the South- ern Ocean utilizing the USNS Eltanin. Samples taken with opening-closing B? plankton nets and Isaacs-Kidd mid- water trawls on these cruises were made available by the Smithsonian Institution for study through the Smithsonian Oceanographic Sorting Center. The exhaustive taxonomic works by Park (1978, 1980, 1982, 1983a, 1983b, 1988, 1993) are based almost exclusively on the midwater trawl samples collected during the Eltanin cruises, and these results signifi cantly increased taxonomic understanding of most species of pelagic calanoids. Other studies based on the Eltanin samples include the following: Bj?rnberg?s (1968) work on the Megacalanidae; Heron and Bowman?s 12_Park_pg143-180_Poles.indd 14412_Park_pg143-180_Poles.indd 144 11/17/08 8:30:12 AM11/17/08 8:30:12 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 145 (1971) on postnaupliar developmental stages of three spe- cies belonging to the genera Clausocalanus and Ctenocala- nus; Bj?rnberg?s (1973) survey of some copepods from the southeastern Pacifi c Ocean; Yamanaka?s (1976) work on the distribution of some Eucalanidae, Aetideidae, and Euchaetidae; Fontaine?s (1988) on the antarctica species group of the genus Paraeuchaeta; Markhaseva?s (2001) on the genus Metridia; and Markhaseva and Ferrari?s (2005) work on Southern Ocean species of Lucicutia. In addition, four exhaustive monographs have treated the taxonomy of a pelagic calanoid family throughout the world?s oceans, and these also have contributed further to an understand- ing of the Southern Ocean fauna: Damkaer?s (1975) work on the Spinocalanidae, Park?s (1995) on the Euchaetidae, Markhaseva?s (1996) on the Aetideidae, and Park?s (2000) on the Heterorhabdidae. These monographs were based on specimens from an extensive set of samples from the world?s oceans. As a result, the taxonomy and geographi- cal range of most of the widespread species of these fami- lies are now well known. Recently described new species of Southern Ocean calanoids are either pelagic species that previous authors failed to recognize as distinct from similar relatives, e.g., Pleuromamma antarctica Steuer, 1931 (see Ferrari and Saltzman, 1998), or species inhabiting extraordinary habitats seldom explored previously, like the water im- mediately above the seafl oor. Bradford and Wells (1983) described the fi rst benthopelagic calanoid copepods of the Southern Ocean, Tharybis magna and Xanthocalanus harpagatus, from a bait bottle. Neoscolecithrix antarctica was collected in small numbers in the Antarctic Sound ad- jacent to the Antarctic Peninsula by Hulsemann (1985b), who believed the species lived in close proximity to the seafl oor. More recent additions to the benthopelagic cala- noid fauna include Parabradyidius angelikae Schulz and Markhaseva, 2000, Paraxantharus brittae Schulz, 2006, and Sensiava longiseta Markhaseva and Schulz, 2006, each belonging to a new genus, and Brachycalanus ant- arcticus Schulz, 2005, Scolecitrichopsis elenae Schulz, 2005, Byrathis arnei Schulz, 2006, Pseudeuchaeta arcu- ticornis Markhaseva and Schulz, 2006, Bradyetes curvi- cornis Markhaseva and Schulz, 2006, Brodskius abyssalis Markhaseva and Schulz, 2007, Rythabis assymmetrica Markhaseva and Schulz, 2007, and Omorius curvispinus Markhaseva and Schulz, 2007. These latter species were collected from the Weddell Sea, an arm of the Southern Ocean, and from the Scotia Sea. In this paper, all relevant studies of the taxonomy of pelagic Southern Ocean calanoid copepods are reviewed. Lists are compiled of species, genera, and families, and the geographical range within the Southern Ocean and rela- tive abundance of each species are noted. Morphological differences are used to suggest evolutionary relationships among species. The distribution of all species is reviewed, and several generalized patterns are hypothesized. Of par- ticular interest here are the species for which fewer than 50 specimens have been collected during the extensive history of surveys of the Southern Ocean pelagic fauna. The distribution of these rare, pelagic calanoids, almost all deepwater species, contributes favorably to an under- standing of patterns of distribution and of speciation in the Southern Ocean. METHODS Traditionally, the Southern Ocean has been described physiographically as including the ocean basins adjacent to the continent of Antarctica plus the following adjoining seas: Amundsen Sea, Bellingshausen Sea, Ross Sea, Wed- dell Sea, and the southern part of the Scotia Sea. In this review of pelagic calanoid copepods, the southern bound- ary of the Southern Ocean is defi ned physiographically by the Antarctic continent, but the northern boundary is defi ned hydrographically, by the average position of the Subtropical Convergence. The Subtropical Convergence is located around 40?S (Deacon 1934, 1937), where the surface temperature of the sea drops sharply from about 18?C to 10?C. In this review, then, the Southern Ocean includes both the Antarctic region and the subantarctic re- gion. Antarctic and subantarctic regions are separated by the Antarctic Convergence, which is located around 55?S, where the sea surface temperature drops 3?C to 5?C over about 30 miles (48.3 km). Although the latitudinal posi- tions of both the Subtropical Convergence, among the At- lantic, Pacifi c and Indian oceans, and the Antarctic Con- vergence, among the Atlantic, Pacifi c and Indian sectors of the Southern Ocean, may vary signifi cantly, locally, these convergences seldom vary more than a degree of latitude from their mean position. Studies of the distribution of organisms are one of the primary purposes of the discipline of taxonomy, and the scope and effectiveness of taxonomic studies is dictated by the availability of specimens. Like most pelagic organisms, calanoid copepods in the Southern Ocean have been col- lected mainly with tow nets operated aboard oceangoing ships, which usually sail to a preselected set of geographic positions in the ocean. Because of the physical isolation of the Southern Ocean, studies of its pelagic calanoid copepods have depended primarily on efforts of national 12_Park_pg143-180_Poles.indd 14512_Park_pg143-180_Poles.indd 145 11/17/08 8:30:13 AM11/17/08 8:30:13 AM 146 SMITHSONIAN AT THE POLES / PARK AND FERRARI oceanographic expeditions, many of which routinely car- ried out sampling protocols for pelagic organisms. The Isaacs-Kidd midwater trawls employed by the U.S. Antarctic Research Program were particularly effec- tive in collecting large, pelagic copepods and signifi cantly increased knowledge about the calanoid fauna. More than 1,000 midwater trawl samples were taken, and these sam- ples are believed to have collected nearly all of the pelagic calanoid copepods in the water column; most of these spe- cies have been described (Park, 1978, 1980, 1982, 1983a, 1983b, 1988, 1993). The trawls were not fi tted with a device to measure water fl ow through the mouth of the trawl, so no quantitative measure of the amount of water fi ltered by the trawl can be calculated for these samples. Sampling times, ranging from one to four hours, have al- lowed a calculation of the number of animals collected per unit time of trawl operation for the more abundant species, e.g., Paraeuchaeta antarctica (see Ferrari and Dojiri, 1987, as Euchaeta antarctica), but this measure is too coarse for the rare species that are the primary focus of this study. The Isaacs-Kidd midwater trawls were quickly lowered to a specifi ed deepest depth, obliquely towed at 3-5 knots to a specifi ed shallowest depth, and then quickly retrieved to the surface again. It is not possible to determine the depth of collection for the specimens captured in a sample with this protocol. Furthermore, normally, only one trawl sam- ple was collected at a station, and therefore, only one depth range was sampled at a particular location. As a result of these constraints, studies based on these samples cannot provide direct information about the vertical distribution of the calanoid species. However, by comparing the presence or absence of a species in samples taken to different greatest depths at different locations here or by calculating a fre- quency of occurrence for each sampled depth range relative to all trawls at that depth range (Yamanaka, 1976), it is possible to make a fi rst-order determination of how deep a trawl has to be towed in order to collect a certain species. Southern Ocean pelagic calanoids are categorized here in several different ways. Species collected in the vicinity of continental or insular land masses are inshore species, while species collected away from continental or insular land masses have been categorized to vertical zones as fol- lows: epipelagic (0? 200m), mesopelagic (200? 1,000m), and bathypelagic (1,000? 4,000). The term ?deepwater? refers to the mesopelagic and bathypelagic zones together. In terms of abundance, species are categorized by the number of specimens collected or known from other ex- peditions: CC, very common (over 100 specimens found); C, common (between 99 and 50 specimens found); R, rare (between 49 and 10); and RR, very rare (less than 10 spec- imens found). These are not counts per sample, but are all specimens known to science. A large number of endemic species are found in the Southern Ocean, and these discoveries are placed within the context of endemicity throughout the world?s oceans for two well-studied families, Euchaetidae and Heteror- habdidae. To facilitate comparisons, four noncontiguous areas of interest were defi ned among the world?s oceans: the Southern Ocean, the Arctic (including adjacent boreal seas of the Atlantic and Pacifi c oceans), the eastern Pacifi c Ocean (along the Pacifi c coasts of the Americas, including the boundary currents), and the Indo-West Pacifi c Ocean (in and around the Malay Archipelago). The latter three were chosen as areas of interest because most of the en- demic species of Euchaetidae and Heterorhabdidae not found in the Southern Ocean occur in one of them. So, for example, the Atlantic Ocean was not considered an area of interest because its endemic species occur mainly toward its northern and southern boundaries, and these endemic species could be included in the Arctic Ocean and Southern Ocean areas, respectively. There are no a priori biological hypotheses that support these utilitarian areas of interest, although endemicity is discussed in the context of high primary and secondary productivity. Three broad feeding categories, herbivory, carnivory, and detritivory, are recognized for many pelagic cala- noids. Food preferences of calanoid copepods have not been studied systematically, but the following publica- tions supported the general feeding categories of these taxa: Mullin?s (1967) work on the herbivory of Calani- dae and Eucalanidae, Yen?s (1983) on the carnivory of Euchaetidae, Ohtsuka et al.?s (1997) on the carnivory of derived Heterorhabdidae, Nishida and Ohtsuka?s (1997) on the detritivory of Scolecitrichidae, and Itoh?s (1970) and Matsuura and Nishida?s (2000) on the carnivory of Augaptilidae. Studies of a few species of the large fam- ily Aetideidae (Robertson and Frost, 1977; Auel, 1999) have suggested that these species may be omnivores, but on the basis of the morphology of their mouthparts, they are considered carnivores here. Feeding is connected to environment through areas of high primary and second- ary productivity, and here high primary and secondary productivity is equated with permanent, annually episodic upwelling areas. These areas of upwelling are associated with western boundary currents adjacent to all continents or are associated with three oceanic bands: trans-Southern Ocean, worldwide equatorial, and boreal Pacifi c (LaFond, 1966; Huber, 2002). 12_Park_pg143-180_Poles.indd 14612_Park_pg143-180_Poles.indd 146 11/17/08 8:30:14 AM11/17/08 8:30:14 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 147 Descriptions of the geographical distribution of a spe- cies beyond the Southern Ocean were initiated by divid- ing the world?s oceans into the following regions, generally following Backus (1986): Antarctic (south of the Antarc- tic Convergence), subantarctic (between the Antarctic and Subtropical convergences), south temperate ( Subtropical Convergence to Tropic of Capricorn, about 20?S), tropi- cal (Tropic of Capricorn to Tropic of Cancer, about 20?N), north temperate (Tropic of Cancer to about 50?N), subarctic (boreal seas adjacent to the Arctic Ocean to the Arctic Cir- cle at 66?N), and the region of the Arctic basin. Some of the boundaries of these areas correspond to hydrographic fea- tures, e.g., they are the surface manifestations of the bound- aries of water masses. However, these boundaries have not been shown to describe the generalized distribution of deep- water animals, and, in fact, two other ways of explaining the distribution of deepwater animals are proposed here. Distribution records for the regions outside of the Southern Ocean are derived from the latest reference given to each species in Appendix 1. There has been no attempt to correct for differences in sampling intensity among these regions, although the south temperate region has been sampled least (only mentioned in reports by Grice and Hulsemann, 1968, and Bj?rnberg, 1973), and the north temperate, subarctic, and Arctic have been sampled the most. Most of the very common pelagic calanoids are usu- ally widespread within the Southern Ocean and were dis- covered during the early expeditions that ended in the fi rst part of the twentieth century. These species originally were described from specimens collected in the Southern Ocean rather than being inferred from descriptions of specimens from other oceans, and most of these species have been re- described once or twice since their original description. As a result, their taxonomy is stable, and their morphology and distribution within the Southern Ocean are relatively well known. Many deepwater calanoids also originally have been described from the Southern Ocean. Their mor- phology is well known, but most of these deepwater species are rare. As a result, information about their distribution, particularly beyond the Southern Ocean, remains limited. Occurrences of these rare deepwater calanoids are based on only a few specimens captured at only a small number of locations, but at least some of the occurrences beyond the Southern Ocean have been confi rmed by direct com- parison of specimens. In contrast, other rare deepwater species originally were described from oceans other than the Southern Ocean and only subsequently were recorded from the Southern Ocean. Because specimens of these rare species from the Southern Ocean have not been compared to specimens from the type locality, their nominal attribu- tion has yet to be verifi ed. In this study, the distribution ranges of many species outside the Southern Ocean are determined from infor- mation available in the literature. Observations about distributions that are accompanied by species descriptions are the usual source for these data. However, not all spe- cies descriptions in the literature are equally informative, and in some cases, it is not easy to determine the identity of all species reported under the same name. In the pres- ent study, the distribution range assigned for many of the rare or very rare species, especially for those reports of a few specimens from several localities, should be consid- ered provisional. The defi nitive answer to the geographi- cal distribution for these rare species must wait until their potential distribution range has been more extensively sampled. For rare and, particularly, very rare species, speci- mens may not have been reported from all of the regions between the Southern Ocean and the region farthest north from which a species has been collected. Initially, a con- tinuous deepwater distribution is assumed if a species is present in more than one nonadjacent region, although a possible origin for disjunct distributions is considered here within a larger context of the evolution of bipolar species pairs. Because the northern boundary of the Southern Ocean is defi ned hydrographically, certain warm-water species have been reported as penetrating for a relatively short distance south into the subantarctic region, while some typically subantarctic species have often been found to extend north of 40?S in small numbers. If the number of these reports is small, an extension of the species range is not considered here. In this study, warm-water species included in the list as subantarctic species are those that have been reported several times, south of the Subtropi- cal Convergence and close to the Antarctic Convergence. Problems have been resolved in a similar manner for range extensions of species between the Antarctic and subant- arctic regions. The identifi cation of a species as having an Antarctic distribution as opposed to a subantarctic distri- bution is based on the number of reports in one water mass relative to the number in the other water mass. In this study, the morphological divergence of the exo- skeleton among species is used as a fi rst approximation to the degree of relatedness among species, with the under- standing that allopatric sibling species may not undergo signifi cant morphological divergence in the absence of a strong adaptive pressure and that secondary sex characters 12_Park_pg143-180_Poles.indd 14712_Park_pg143-180_Poles.indd 147 11/17/08 8:30:14 AM11/17/08 8:30:14 AM 148 SMITHSONIAN AT THE POLES / PARK AND FERRARI of the exoskeleton are likely to diverge over a shorter period of time than the rest of the exoskeleton. Specimens from the wider geographical regions are reexamined for information regarding the morphological variation and distribution ranges of some of the species (T. Park, unpub- lished data). The abundances found in Tables 1? 7 are derived from Park (1978, 1980, 1982, 1983a, 1983b, 1988, 1993), who examined samples selected mainly from the Eltanin mid - water trawl collections available from the Smithsonian Oceanographic Sorting Center. The selection was made to cover as wide an area as possible, but only an aliquot of the original sample was examined systemically and consis- tently so that the sorted specimens refl ect the relative abun- dance of the species in the sample in a general way. In this paper?s introduction, counts of the number of species of co- pepods and of calanoid copepods were determined by using an online species database (The World of Copepods, Na- tional Museum of Natural History) to extend the counts of Humes (1994), beginning with the year 1993, and Bowman and Abele (1982), beginning with the year 1982. NUMBER SPECIES OF PELAGIC CALANOIDS FROM THE SOUTHERN OCEAN Two hundred and fi ve species of pelagic calanoid cope- pods (Appendix 1) in 57 genera from 21 families (Appendix 2) have been reported from the Southern Ocean. Among these, 8 (3.9%) species are coastal or inshore, 13 (6.3%) are epipelagic species, and 184 (89.8%) are deepwater species. Of the 57 genera of pelagic calanoids reported from the Southern Ocean, Paraeuchaeta is the most speciose genus with 21 species, followed by Scolecithricella (16 species), Euaugaptilus (14 species), Scaphocalanus (10 species), Metridia (9 species), Pseudochirella (9 species), Gaetanus (8 species), Onchocalanus (7 species), and Lucicutia (6 spe- cies). The 16 species included in the genus Scolecithricella are morphologically diverse, and recently, Vyshkvartzeva (2000) proposed to place the species of Scolecithricella in one of three genera. Although we are not comfortable with this proposal because the analysis was not exhaustive, ac- ceptance would result in the following numbers from the Southern Ocean: Scolecithricella (9 species) and Pseu- doamallothrix (6 species), with S. pseudopropinqua being moved to the genus Amallothrix. Species of Paraeuchaeta (Euchaetidae) are deepwater calanoids, and some of them can be collected in large num- bers in the Southern Ocean, e.g., Paraeuchaeta antarctica (see Park, 1978; Ferrari and Dojiri, 1987) or P. barbata, P. rasa, and P. biloba (see Park, 1978). Feeding of a few species of Paraeuchaeta has been studied (Yen, 1983, 1991), and these species are known to be carnivores. On the basis of the similarity of feeding appendages among species, this feeding mode is assumed for all species of the genus. In the Southern Ocean, species of Euaugapti- lus (Augaptilidae) are typically bathypelagic and thought to be carnivores on the basis of the structure of their feeding appendages (Itoh, 1970; Matsuura and Nishida, 2000); they have not been reported in large numbers. The family Aetideidae has the largest number of species (45) in the Southern Ocean, with almost 40% of its species belonging to two genera, Pseudochirella and Gaetanus; these species also are considered to be carnivores. The combined 80 species of Paraeuchaeta, Euaugaptilus, and Aetideidae, then, represent a signifi cant contribution to carnivory in the Southern Ocean and so are believed to play a major role in the dynamics of the deepwater plank- ton community. The Scolecitrichidae is the next most speciose family after Aetideidae. Among the 35 species of Scolecitrichi- dae, those in the genera Scolecithricella and Scaphocala- nus make up 45% of the family in the Southern Ocean. Although the feeding niche of most species of Scolecit- richidae is not well known, derived chemosensory setae on maxilla 2 and maxilliped (Nishida and Ohtsuka, 1997) suggest that species of Scolecitrichidae are the major de- tritivores in the Southern Ocean. In contrast to carnivory and detritivory, herbivory in the Southern Ocean has been studied more extensively (Hardy and Gunther, 1935; An- drews, 1966). The herbivore fauna is structured by large numbers of individuals belonging to a few species in gen- era of two families: Calanus (three species) and Calanoi- des (one species) in Calanidae and Eucalanus (one species) and Rhincalanus (one species) in Eucalanidae. Phylogenetic relationships among the congeneric spe- cies of the Southern Ocean have been proposed for two families that have been studied worldwide, Euchaetididae and Heterorhabdidae. Six monophyletic groups of spe- cies (species groups) of Paraeuchaeta have been identifi ed (Park, 1995). One or more species in fi ve of the six species groups of Paraeuchaeta are found in the Southern Ocean. One species group, the antarctica species group, consists of fi ve species, and all of its species are limited to waters south of the Antarctic Convergence; this is the only spe- cies group of either family that is found only in the South- ern Ocean. Three species in the antarctica species group of Paraeuchaeta appear to be restricted to waters along the edge of ice shelf of Antarctica, and all fi ve may share 12_Park_pg143-180_Poles.indd 14812_Park_pg143-180_Poles.indd 148 11/17/08 8:30:15 AM11/17/08 8:30:15 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 149 this same inshore habitat (Fontaine, 1988). Species of the antarctica species group are assumed to have evolved after the colonization of the Southern Ocean by a com- mon ancestor (see the Evolution of the Pelagic Calanoid Fauna within the Southern Ocean section). Each species of Paraeuchaeta in the remaining four species groups rep- resented in the Southern Ocean has its closest relative in other oceans, rather than the Southern Ocean, suggesting that the remaining species of Paraeuchaeta may have colo- nized the Antarctic region independently. The family Heterorhabdidae is represented in the Southern Ocean by three genera, Heterorhabdus, Hetero- stylites, and Paraheterorhabdus. There are fi ve species in the fi rst genus; a single species in each of the last two genera is found in the Southern Ocean (Park, 2000). Four of the fi ve species of Heterorhabdus belong to the same species group, the abyssalis species group, with 17 species. Two of the four species of this group, H. spinosus and H. paraspi- nosus, are morphologically quite similar, suggesting a recent speciation event within the Southern Ocean. The remaining two Heterorhabdus species of the abyssalis species group, H. austrinus and H. pustulifer, are morphologically dis- similar; they may have colonized the Southern Ocean inde- pendently from one another and from the pair H. spinosus and H. paraspinosus. The fi fth species, H. lobatus, belongs to the papilliger species group along with fi ve other species found in other oceans; along with Paraheterorhabdus far- rani and Heterostylites nigrotinctus, H. lobatus represents an independent colonization. INSHORE CALANOIDS ALONG CONTINENTAL AND INSULAR COASTS OF THE SOUTHERN OCEAN Eight species of pelagic calanoids have been found exclusively in waters close to a land mass of the Southern Ocean (Table 1): three species of Drepanopus (D. bispi- nosus, D. forcipatus, and D. pectinatus), three species of Paralabidocera (P. antarctica, P. grandispina, and P. separabilis), and two species of Stephos (S. longipes and S. antarcticus). The three species of Drepanopus have been collected several times, and all three occa- sionally have been collected in very large numbers, so their distribution is well known. Drepanopus bispinosus has been reported, often as abundant, from inshore wa- ters adjacent to the Vestfold Hills region of Antarctica (Bayly, 1982), and its population structure has been es- tablished (Bayly, 1986). Drepanopus pectinatus occurs close to shores of Crozet Island, Kerguelen Island, and Heard Island in the Indian Ocean sector of the Southern Ocean (Hulsemann, 1985a); some aspects of its biology also have been elucidated (Razouls and Razouls, 1990). Drepanopus forcipatus is restricted to Atlantic and Pa- cifi c coastal and shelf areas along southern South Amer- ica, including the Falkland Islands, and around South Georgia Island (Hulsemann, 1985a); its copepodid stages have been described (Hulsemann, 1991). The distributions of the remaining fi ve inshore species are not as well known, and only Paralabidocera antarctica and Stephos longipes have been reported from more than one locality. Paralabidocera antarctica is now known to occur in small numbers in waters close to the shoreline at several locations, including the South Shetland Islands, the extreme south of the Ross Sea, two localities in the At- lantic Ocean sector of Antarctica, and one locality in the Indian Ocean sector of Antarctica (Vervoort, 1957). The species is believed to inhabit the surface water layers and is occasionally captured under the ice (Vervoort, 1951); development of its marine and lacustrine populations has been described (Swadling et al., 2004). Paralabidocera grandispina Waghorn, 1979 and P. separabilis Brodsky and Zvereva, 1976 are known only from their type lo- calities, beneath the ice along the Pacifi c Ocean sector of Antarctica and near the shore of Antarctica in the Indian Ocean sector, respectively. Of the two species of Stephos, S. longipes has been found close to or under the ice shelf in the Pacifi c and Indian ocean sectors of Antarctica including the Ross Sea, where it can be very abundant (Giesbrecht, 1902; Farran, 1929; Tanaka, 1960). Its associations with the ice and the open water have been described (Kurbjeweit et al., 1993; TABLE 1. Inshore pelagic calanoid copepods of the southern ocean. Ant, waters south of the Antarctic Convergence; S-Ant, waters between the Antarctic Convergence and the Subantarctic Convergence; CC, very common (over 100 specimens found); C, common (between 99 and 50 specimens found); R, rare (between 49 and 10 specimens found). Species name Distribution Abundance Drepanopus bispinosus Ant CC Drepanopus forcipatus S-Ant CC Drepanopus pectinatus S-Ant CC Paralabidocera antarctica Ant C Paralabidocera grandispina Ant R Paralabidocera separabilis Ant R Stephos longipes Ant C Stephos antarcticus Ant R 12_Park_pg143-180_Poles.indd 14912_Park_pg143-180_Poles.indd 149 11/17/08 8:30:15 AM11/17/08 8:30:15 AM 150 SMITHSONIAN AT THE POLES / PARK AND FERRARI Schnack-Schiel et al., 1995). Stephos antarcticus is known only from its type locality, McMurdo Sound of the Ross Sea (Wolfenden, 1908). Most species of the genus Stephos are closely associated with the water immediately above the seafl oor (Bradford-Grieve, 1999). Among these eight species, the three species of Drepa- nopus and the three species of Paralabidocera clearly seem to be pelagic. The two species of Stephos appear to be ice oriented but are considered pelagic here. All of these inshore species are characteristically small in size, ranging from 0.85 to 2.80 mm in body length. EPIPELAGIC FAUNA OF THE SOUTHERN OCEAN The epipelagic calanoid fauna of the Southern Ocean south of the Antarctic Convergence (Table 2) is relatively simple in species composition. There are fi ve species, all are very common, and their combined biomass is unsurpassed by the epipelagic calanoid fauna of any other region of the world?s oceans (Foxton, 1956). These fi ve epipelagic spe- cies are, in order of abundance, Calanoides acutus, Rhin- calanus gigas, Calanus propinquus, Metridia gerlachei, and Clausocalanus laticeps. These are the copepods most often associated by planktonologists with the Southern Ocean. All were discovered during the early expeditions, and their taxonomy and distribution have been clearly and carefully defi ned. Although these fi ve species are more abundant in waters south of the Antarctic Convergence, they also may be collected north of the convergence, but here they appear to be associated with the deeper Antarc- tic Intermediate Water. Calanoides acutus and, to a lesser extent, Rhincalanus gigas and Calanus propinquus are the dominant herbivores south of the Antarctic Convergence, and their role in that ecosystem is well known (Chiba et al., 2002; Pasternak and Schnack-Schiel, 2001). The analogous epipelagic calanoids of the subantarc- tic region, between the Antarctic and Subtropical conver- gences, are Calanus simillimus, Clausocalanus brevipes, and Ctenocalanus citer. These three herbivores are very common in the subantarctic region, but they are not as numerous in these waters as the previous fi ve epipelagic calanoids are south of the Antarctic Convergence. Fur- thermore, the Antarctic Convergence does not limit the southern boundary of the range of these three species as precisely as it limits the northern boundary of the previous fi ve epipelagic calanoids. The population structure and life histories of the three have been described (Atkinson, 1991; Schnack-Schiel and Mizdalski, 1994). There are three additional large-sized, epipelagic her- bivores that may be collected in the subantarctic region as well as in the south temperate midlatitudes: Calanus australis, Neocalanus tonsus, and Subeucalanus longiceps. Calanus australis is known to be distributed along the southern coast of Chile, off Argentina, in New Zealand waters, and in southeastern Australian waters (Bradford- Grieve, 1994). Its distribution during summer has been investigated (Sabatini et al., 2000). Neocalanus tonsus is widely distributed in subantarctic waters but also may be found in the deepwater of the south temperate region; some aspects of its life history are known (Ohman et al., 1989). Subeucalanus longiceps (Subeucalanidae) occurs circumglobally in the subantarctic and temperate regions of the Southern Hemisphere. These three species are im- portant herbivores in the subantarctic as well as the south temperate region. Two small-sized, epipelagic herbivores, Clausocalanus parapergens and Ctenocalanus vanus, are found in sub- antarctic waters. Clausocalanus parapergens has been re- ported as far north as the subtropical convergence (Frost and Fleminger, 1968). Specimens referred to as Ctenocala- nus vanus from the Southern Ocean by Farran (1929) and Vervoort (1951, 1957) are Ctenocalanus citer (T. Park, TABLE 2. Epipelagic calanoid copepods of the Southern Ocean. CC, very common (over 100 specimens found); C, common ( between 99 and 50 specimens found). Species name Abundance Species endemic to Antarctic waters Calanoides acutus CC Calanus propinquus CC Clausocalanus laticeps CC Metridia gerlachei CC Rhincalanus gigas CC Species endemic to subantarctic waters Calanus simillimus CC Clausocalanus brevipes CC Ctenocalanus citer C Species ranging from subantarctic water to south temperate region Calanus australis C Neocalanus tonsus C Subeucalanus longiceps C Species ranging from subantarctic waters to north temperate region Eucalanus hyalinus C Rhincalanus nasutus C 12_Park_pg143-180_Poles.indd 15012_Park_pg143-180_Poles.indd 150 11/17/08 8:30:16 AM11/17/08 8:30:16 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 151 unpublished observations); this species is restricted to the Southern Ocean. Calanids like Calanoides acutus, Calanus propinquus, Calanus simillimus, Calanus australis, and Neocalanus tonsus as well as eucalanids like Rhincalanus gigas and subeucalanids like Subeucalanus longiceps are consid- ered epipelagic here because they spend their juvenile and adult life in near-surface waters. However, during seasonal episodes of low primary productivity, some late juvenile stages of the populations of each of these species descend to mesopelagic depths (Vervoort, 1957) to diapause. Two epipelagic calanoid species, Eucalanus hyalinus and Rhincalanus nasutus, do not occur in large numbers in the subantarctic region. North of the Subtropical Con- vergence, they often are encountered in warmer waters, and they have been collected in the north temperate re- gion. Their taxonomy and distribution are well under- stood. Usually, only a few specimens are collected in pe- lagic samples from the Southern Ocean, and here these species are considered to be associated with habitats of low primary productivity. In summary, 13 calanoid species contribute to the epipelagic fauna of the Southern Ocean. Five of them are endemic south of the Antarctic Convergence and are very common throughout this region. Three species are endemic to the subantarctic region and occur throughout that region. However, the subantarctic endemics are not as numerous as the fi ve species endemic to the Antarctic region in midwater trawl samples. Three more epipelagic calanoid species occur widely in the subantarctic and temperate regions of the Southern Hemisphere. They are either common in productive coastal upwelling areas or are circumglobal in the West Wind Drift of the Southern Hemisphere. The broadest latitudinal range exhibited by subantarctic epipelagic calanoids is that of the two species of Eucalanidae that have been collected from the subant- arctic region to the north temperate region. DEEPWATER CALANOIDS RESTRICTED TO THE SOUTHERN OCEAN Among the 184 species of deepwater calanoids found in the Southern Ocean, 50 species were originally described from the Southern Ocean and, to date, are known exclu- sively from there (Table 3). Twenty-four of these deepwa- ter calanoids originally were described from waters south of the Antarctic Convergence and subsequently have been found exclusively in those waters; they are endemics of the Antarctic region. Of these 24 Antarctic endemics, six species have a distinctly localized distribution, occurring almost exclusively along the ice edge of Antarctica. There are four closely related species of the antarctica species group of Paraeuchaeta, plus one species each of Aetideop- sis and Chiridiella. All six species have strongly built bod- ies and limbs and a well-sclerotized exoskeleton; they are presumed to be carnivores. Chiridiella megadactyla was described from a single female collected close to the edge of the Ross Ice Shelf and has not been found again. Aetide- opsis antarctica, a rare species, was collected initially from waters beneath the edge of the Ross Ice Shelf (Wolfenden, 1908); it subsequently has been found several other times from the same habitat. Three of the four species of the ant- arctica species group of Paraeuchaeta, P. austrina, P. erebi, and P. tycodesma, have also been reported only a few times and collected only in small numbers. Paraeuchaeta similis, the fourth species of the group, has occasionally been reported to be quite common under the ice (Brad- ford, 1981), unlike the above three congeners of its spe- cies group, which are rare. However, P. similis also may be collected in the deeper layer of warm water (Vervoort, 1965b), well away from the ice edge. Here it occasionally may co-occur with P. antarctica (see Ferrari and Dojiri, 1987), the fi fth species of the group (Fontaine, 1988; Park, 1995). The three ice edge species of Paraeuchaeta, together with P. similis and P. antarctica, a species endemic to and abundant throughout Antarctic and the subantarctic re- gions (Park, 1978; Mar?n and Antezana, 1985; Ferrari and Dojiri, 1987), form the group of closely related species. Despite this close relationship, all fi ve of these species have been collected a number of times in the same midwater trawl from waters adjacent to the ice edge. Four other species, Batheuchaeta antarctica, B. pube- scens, Pseudochirella formosa, and Onchocalanus subcris- tatus, were initially described from deep water south of the Antarctic Convergence. All have been collected only once, and each is known only from one or two specimens. The fi rst three are aetideids and are presumed to be car- nivores; the last belongs to Phaennidae, a family of detri- tivores related to the Scolecitrichidae. These four species and the earlier mentioned Chiridiella megadactyla remain so poorly known that their taxonomic status and distribu- tion cannot be confi rmed. Among the 24 species endemic to the Antarctic region, the remaining 14 species have been collected widely south of the Antarctic Convergence and, except for Euaugapti- lus austrinus and Landrumius antarcticus, are either com- mon or very common, so their taxonomy and distribution have been well established. Among them are three species of small calanoid copepods, Scaphocalanus vervoorti, 12_Park_pg143-180_Poles.indd 15112_Park_pg143-180_Poles.indd 151 11/17/08 8:30:16 AM11/17/08 8:30:16 AM 152 SMITHSONIAN AT THE POLES / PARK AND FERRARI Scolecithricella cenotelis, and Scaphocalanus subbrevicor- nis, that may occur in particularly large numbers in wa- ters close to continent, where they may be encountered in relatively shallow water (Park, 1980, 1982). These small, abundant species all belong to the family Scolecitrichidae and are presumed to be detritivores. Two other small, common, pelagic calanoids, Scolecithricella vervoorti and Spinocalanus terranovae, are found exclusively in the Ant- arctic region but in relatively smaller numbers than the fi rst three. The former is a scolecitrichid. Spinocalanus ter- ranovae belongs to the Spinocalanidae; its trophic niche is not known. Of the remaining nine deepwater species restricted to waters south of the Antarctic Convergence, all are relatively large calanoids. They can be divided into two groups. Four species are very common; in order of the number of speci- mens collected they are Euchirella rostromagna, Halopti- lus ocellatus, Scaphocalanus antarcticus, and Euaugapti- lus antarcticus. Three species are common, Onchocalanus wolfendeni, Paraeuchaeta eltaninae, and Onchocalanus TABLE 3. Abundances of deepwater calanoid species endemic to the Southern Ocean. CC, very common; C, common; R, rare; RR, very rare. Species name Abundance Species name Abundance Species occurring along the ice edge of Antarctica (6 spp.) Aetideopsis antarctica R Chiridiella megadactyla Paraeuchaeta austrina Paraeuchaeta erebi Paraeuchaeta similis C Paraeuchaeta tycodesma R Species occurring widely in Antarctic waters (14 spp.) Euaugaptilus antarcticus CC Euaugaptilus austrinus Euchirella rostromagna CC Haloptilus ocellatus Landrumius antarcticus R Onchocalanus magnus C Onchocalanus wolfendeni Paraeuchaeta eltaninae Scaphocalanus antarcticus CC Scaphocalanus subbrevicornis Scaphocalanus vervoorti CC Scolecithricella cenotelis Scolecithricella vervoorti C Spinocalanus terranovae Species known from 1 or 2 specimens in Antarctic waters (4 spp.) Batheuchaeta antarctica RR Batheuchaeta pubescens Onchocalanus subcristatus RR Pseudochirella formosa Species occurring in both Antarctic and subantarctic waters (19 spp.) Aetideus australis C Candacia maxima R Cephalophanes frigidus Heterorhabdus pustulifer Heterorhabdus austrinus Heterostylites nigrotinctus Metridia pseudoasymmetrica R Paraeuchaeta antarctica CC Paraeuchaeta biloba Paraeuchaeta dactylifera C Paraeuchaeta parvula Paraeuchaeta rasa CC Paraheterorhabdus farrani Pleuromamma antarctica Pseudochirella mawsoni C Scaphocalanus farrani CC Scaphocalanus parantarcticus CC Scolecithricella dentipes CC Scolecithricella schizosoma Species endemic to subantarctic waters (7 spp.) Aetideopsis tumorosa R Bathycalanus eltaninae Bathycalanus unicornis Bradycalanus enormis Bathycalanus infl atus Bradycalanus pseudotypicus R Candacia cheirura C magnus, and two species are rare, Euaugaptilus austri- nus and Landrumius antarcticus. These large species are taxonomically diverse and belong to fi ve calanoid families (Appendix 2). There are 19 endemic species of calanoid copepods that have been found in both the Antarctic and the sub- antarctic regions, i.e., south of the Subtropical Conver- gence. Most prominent among them are fi ve species of Paraeuchaeta: P. antarctica, P. biloba, P. rasa, P. parvula, and P. dactylifera. Paraeuchaeta antarctica and P. rasa are among the most abundant carnivorous calanoids of the Southern Ocean. They are usually encountered south of the Antarctic Convergence but may be collected in small numbers to the north in open waters; P. antarctica has also been reported as far north as the Chilean fjords (Mar?n and Antezana, 1985). Paraeuchaeta biloba can be collected immediately adjacent to, and on either side of, the Antarctic Convergence. A unique record of the co- occurrence of these three species of Paraeuchaeta is from a deep midwater trawl sample (0? 1,295 m) taken 12_Park_pg143-180_Poles.indd 15212_Park_pg143-180_Poles.indd 152 11/17/08 8:30:17 AM11/17/08 8:30:17 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 153 off Uruguay (34?43H11032S, 49?28H11032W to 34?51H11032S, 49?44H11032W) in the southwestern Atlantic north of the Subtropical Con- vergence (Park, 1978). Paraeuchaeta parvula, like P. biloba, has also been collected both north and south of the Antarctic Conver- gence. Paraeuchaeta dactylifera has only been found in relatively small numbers and usually in the subantarctic region; two specimens captured well south of the Antarc- tic Convergence (Park, 1978) are exceptions. Two aetideid species, Aetideus australis and Pseudochirella mawsoni, can also be found both north and south of the Antarctic Convergence. Aetideus australis has been collected more often in waters north of the convergence than south. Pseu- dochirella mawsoni has been reported from numerous lo- calities in the Southern Ocean and has been collected in large numbers in midwater trawls immediately north of convergence. Both species are presumed to be carnivores. Four very common scolecitrichid species are endemic to the Southern Ocean. Scolecithricella dentipes and Scaphocalanus farrani are found throughout the Antarctic and subantarctic regions, where they may be numerous in some samples. Scaphocalanus parantarcticus and Scole- cithricella schizosoma are also distributed throughout the Southern Ocean and may be very common but are found in smaller numbers than the fi rst two. There are four species of the family Heterorhabdidae that are well-known endemics of the Southern Ocean: Het- erorhabdus pustulifer, H. austrinus, Heterostylites nigro- tinctus, and Paraheterorhabdus farrani. Paraheterorhabdus farrani is common and has been collected throughout the Southern Ocean; Heterorhabdus pustulifer and H. austri- nus are also common, while Heterostylites nigrotinctus is rare. Among the remaining 4 of the 19 endemic species re- ported from both Antarctic and subantarctic regions, three rare species, Candacia maxima, Cephalophanes frigidus, and Metridia pseudoasymmetrica, and the common Pleu- romamma antarctica are not often encountered in samples. However, there are enough records to suggest that these species are limited to the Southern Ocean. There are seven deepwater species that have been found only in the subantarctic region (Table 3). Six of them, Aetideopsis tumerosa, Bathycalanus eltaninae, B. unicornis, Bradycalanus enormis, B. infl atus, and B. pseudotypicus, are known from a few localities and only a few specimens; their distribution cannot be determined with certainty. Of these six species, the latter fi ve belong to the family Megacalanidae; Aetideopsis tumerosa is an aetideid. The seventh species, Candacia cheirura, has been collected often enough to be considered the only species of Candaciidae restricted to the subantarctic region. It is common and has been hypothesized to be restricted to me- sopelagic waters of the West Wind Drift (Vervoort, 1957), also called the Antarctic Circumpolar Current, which is the dominant circulation feature of the Southern Ocean. In summary, among the 50 deepwater calanoid cope- pod species found exclusively in the Southern Ocean, six species occur close to the continent. Although these species have been captured in relatively small numbers, they may have been undersampled due to the diffi culty in collecting with a midwater trawl in deepwater close to the continent. Among these six species, the closely related Paraeuchaeta austrina, P. erebi, and P. tycodesma, all members of the antarctica species group, appear to be restricted to the same habitat. Of the 18 species found in open waters south of the Antarctic Convergence, four were originally described from one or two specimens collected in a single sample, have not been rediscovered, and remain poorly known. The remaining 14 species can be regarded as typi- cal endemics of the Antarctic deep water. Except for two relatively rare species, they are common or very common in waters south of the Antarctic Convergence. Nineteen of the 50 Southern Ocean deepwater species are typical endemics of the region as a whole, and most of them have been collected from many localities throughout the South- ern Ocean. Seven of these species have only been found in the subantarctic region. Their distributions are based on a small number of specimens and therefore are insuffi ciently known. Candacia cheirura is an exception; it is a common endemic of the subantarctic region. DEEPWATER CALANOIDS FROM ANTARCTIC WATERS REPORTED NORTH OF THE SUBTROPICAL CONVERGENCE A total of 127 deepwater species of pelagic cala- noid copepods have been reported from the Southern Ocean south of the Antarctic Convergence. Twenty-four of those species are limited to this region (see the Deep- water Calanoids Restricted to the Southern Ocean sec- tion), and 19 species have been found northward, into the subantarctic region, with their distribution terminat- ing at the Subtropical Convergence. Thus, 43 of these 127 deep water species collected south of the Antarctic Convergence are endemic to the Southern Ocean. The remaining 84 species have been reported beyond the Subtropical Convergence to varying degrees. Seven (8%) of these species have been collected in the south tem- perate region, adjacent to the Southern Ocean (Table 4), and fi ve (6%) species have been collected as far north as 12_Park_pg143-180_Poles.indd 15312_Park_pg143-180_Poles.indd 153 11/17/08 8:30:17 AM11/17/08 8:30:17 AM 154 SMITHSONIAN AT THE POLES / PARK AND FERRARI TABLE 4. Abundances and locations of deepwater calanoid species collected south of the Antarctic Convergence that occur north of the Sub tropical Convergence. 1, south temperate; 2, tropical; 3, north temperate; 4, subarctic; 5, Arctic basin. Abundance in Southern Ocean: CC, very common; C, common; R, rare; RR, very rare. A ? H11001 ? indicates presence. Region Species name Abundance 1 2 3 4 5 Species ranging from Antarctic waters to the south temperate region (7 spp.) Euaugaptilus hadrocephalus RR H11001 Euaugaptilus perasetosus R H11001 Onchocalanus paratrigoniceps R H11001 Paraeuchaeta regalis CC H11001 Pseudochirella hirsuta C H11001 Scolecithricella hadrosoma R H11001 Scolecithricella parafalcifer R H11001 Species ranging from Antarctic waters to tropical region (5 spp.) Cornucalanus robustus CC H11001 H11001 Farrania frigida R H11001 H11001 Lucicutia bradyana R H11001 H11001 Paraeuchaeta abbreviata R H11001 H11001 Scaphocalanus major R H11001 Species ranging from Antarctic waters to north temperate region (29 spp.) Batheuchaeta lamellata R H11001 H11001 H11001 Batheuchaeta peculiaris R H11001 H11001 Bathycalanus bradyi R H11001 H11001 H11001 Chiridiella subaequalis R H11001 Chiridius polaris R H11001 Cornucalanus chelifer CC H11001 H11001 H11001 Cornucalanus simplex R H11001 H11001 H11001 Euaugaptilus bullifer R H11001 H11001 Euaugaptilus magna C H11001 H11001 H11001 Euaugaptilus maxillaris R H11001 H11001 H11001 Euaugaptilus nodifrons C H11001 H11001 H11001 Gaetanus antarcticus R H11001 H11001 H11001 Gaetanus paracurvicornis R H11001 H11001 Haloptilus fons R H11001 H11001 H11001 Haloptilus oxycephalus CC H11001 H11001 Lophothrix humilifrons R H11001 H11001 H11001 Metridia ferrarii R H11001 H11001 H11001 Onchocalanus cristatus R H11001 H11001 H11001 Onchocalanus hirtipes R H11001 Onchocalanus trigoniceps R H11001 H11001 H11001 Scaphocalanus elongatus C H11001 H11001 Scolecithricella altera R H11001 Scolecithricella emarginata CC H11001 H11001 H11001 Scolecithricella obtusifrons R H11001 H11001 Scolecithricella ovata C H11001 H11001 H11001 T alacalanus greeni R H11001 H11001 V aldiviella oligarthra R H11001 H11001 H11001 V aldiviella brevicornis R H11001 H11001 H11001 V aldiviella insignis R H11001 H11001 H11001 (continued) 12_Park_pg143-180_Poles.indd 15412_Park_pg143-180_Poles.indd 154 11/17/08 8:30:18 AM11/17/08 8:30:18 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 155 Region Species name Abundance 1 2 3 4 5 Species ranging from Antarctic waters to the subarctic region (30 spp.) Aetideopsis multiserrata R H11001 H11001 H11001 H11001 Arietellus simplex R H11001 H11001 H11001 H11001 Candacia falcifera R H11001 H11001 H11001 Chiridius gracilis R H11001 H11001 H11001 Haloptilus longicirrus R H11001 H11001 H11001 Lucicutia curta R H11001 H11001 H11001 H11001 Lucicutia macrocera R H11001 H11001 H11001 H11001 Lucicutia magna R H11001 H11001 H11001 H11001 Lucicutia ovalis R H11001 H11001 H11001 H11001 Lucicutia wolfendeni R H11001 H11001 H11001 H11001 Megacalanus princeps R H11001 H11001 H11001 H11001 Metridia curticauda R H11001 H11001 H11001 H11001 Metridia ornata R H11001 H11001 Metridia princeps R H11001 H11001 H11001 H11001 Mimocalanus cultrifer R H11001 H11001 H11001 H11001 Nullosetigera bidentatus R H11001 H11001 H11001 H11001 Pachyptilus eurygnathus R H11001 H11001 H11001 H11001 Paraeuchaeta kurilensis R H11001 H11001 H11001 H11001 Paraeuchaeta tumidula R H11001 H11001 H11001 Pseudeuchaeta brevicauda R H11001 H11001 H11001 H11001 Pseudochirella dubia R H11001 H11001 H11001 H11001 Pseudochirella notacantha R H11001 H11001 H11001 Pseudochirella obtusa C H11001 H11001 H11001 Pseudochirella pustulifera R H11001 H11001 H11001 H11001 Racovitzanus antarcticus CC H11001 Scolecithricella minor CC H11001 H11001 H11001 H11001 Scolecithricella valida C H11001 H11001 H11001 H11001 Spinocalanus abyssalis R H11001 H11001 H11001 H11001 Spinocalanus magnus R H11001 H11001 H11001 H11001 Undeuchaeta incisa R H11001 H11001 H11001 Species ranging from Antarctic waters to the Arctic Ocean (13 spp.) Aetideopsis minor R H11001 H11001 Aetideopsis rostrata R H11001 H11001 Augaptilus glacialis R H11001 H11001 H11001 H11001 H11001 Gaetanus brevispinus CC H11001 H11001 H11001 H11001 H11001 Gaetanus tenuispinus CC H11001 H11001 H11001 H11001 H11001 Microcalanus pygmaeus R H11001 H11001 H11001 H11001 H11001 Paraeuchaeta barbata CC H11001 H11001 H11001 H11001 H11001 Pseudaugaptilus longiremis R H11001 H11001 H11001 Pseudochirella batillipa R H11001 H11001 H11001 Pseudochirella spectabilis R H11001 H11001 Spinocalanus antarcticus R H11001 Spinocalanus horridus R H11001 H11001 H11001 H11001 H11001 T emorites brevis R H11001 H11001 H11001 H11001 12_Park_pg143-180_Poles.indd 15512_Park_pg143-180_Poles.indd 155 11/17/08 8:30:18 AM11/17/08 8:30:18 AM 156 SMITHSONIAN AT THE POLES / PARK AND FERRARI the tropical region. There are records of 29 species (35%) from the Southern Ocean as far north as the north temper- ate region and reports of 30 species (36%) as far north as the subarctic seas (Table 5). Thirteen species (15%) have been collected as far north as the Arctic Ocean. Five (71%) of the seven species ranging from the Ant- arctic region to the south temperate region are rare or very rare (Table 4). Euaugaptilus hadrocephalus, E. peraseto- sus, Onchocalanus paratrigoniceps, Scolecithricella hadro- soma, and S. parafalcifer originally were described from a few specimens and have not been collected again; they are poorly known. Two species, Paraeuchaeta regalis and Pseu- dochirella hirsuta, have been collected from many localities throughout the subantarctic region, and specimens have been found in small numbers from samples both northward in the south temperate region and southward into Antarctic regions. Paraeuchaeta regalis, like its euchaetid congeners, is probably a carnivore; Pseudochirella hirsuta, an aetideid, is also presumed to be carnivorous on the basis of the size and structure of its feeding appendages. Of the fi ve Southern Ocean species that have been collected into the deepwater of the tropical region (Table 4), Farrania frigida, Lucicutia bradyana, Paraeuchaeta abbreviata, and Scaphocalanus major are rare (80%) and remain poorly known; their records are based on a small number of specimens collected from a few widely separated localities. Only one of the fi ve species, Cornu- calanus robustus, occurs throughout the Southern Ocean; Park (1983b) recovered it from 37 deepwater stations in the Antarctic and subantarctic regions. Vervoort (1965a) identifi ed fi ve specimens, including two juvenile copepo- dids in the deep water of the Gulf of Guinea in the tropical Atlantic, and this remained the only record from outside the Southern Ocean until Bj?rnberg (1973) reported it in the southeastern Pacifi c Ocean. All 29 Southern Ocean species collected as far north as the north temperate region (Table 4) appear to be bathy- pelagic, found between 1,000 and 4,000 m. Twenty-two (76%) of these are rare in the Southern Ocean. Among the remaining seven species, six are either common or very common (number in the parenthesis is the number of speci- mens found in the Southern Ocean): Haloptilus oxycepha- lus (388), Scolecithricella emarginata (226), Cornucalanus chelifer (111), Scolecithricella ovata (76), Euaugaptilus nodifrons (71), and Scaphocalanus elongatus (63). Scole- cithricella emarginata and Cornucalanus chelifer are very common in the Southern Ocean. Haloptilus oxycephalus is very common and Euaugaptilus nodifrons is common in the Southern Ocean, but only a few specimens of either spe- cies have been collected in the subantarctic, south temper- ate, or tropical regions. Scolecithricella ovata and Scapho- calanus elongatus are common in the subantarctic region. Forty-four specimens of the seventh species, Euaugaptilus magnus, have been recovered from the Southern Ocean, but most of these are from the subantarctic region. It is cat- egorized here as rare but is still better represented than the other 22 rare species. Thirty species reported from Antarctic region also have been collected in the subarctic region (Table 4). All of them are bathypelagic, and 87% are rare. The remaining four species are common or very common in the Southern Ocean (number in the parenthesis is the number of speci- mens found in the Southern Ocean by T. Park): Scolecithri- cella minor (1,728), Racovitzanus antarcticus (1,077), Sco- lecithricella valida (74), and Pseudochirella obtusa (52). A greater number of specimens of Scolecithricella minor than any other species of this genus was encountered in the South- ern Ocean. Specimens were more likely to be collected in water close to the continent, where the species occasionally has been reported from the epipelagic zone. Racovitzanus antarcticus is more likely to be encountered in the Antarctic region although it also occurs in waters immediately north of the Antarctic Convergence and beyond. Scolecithricella valida was found widely throughout the Southern Ocean. Pseudochirella obtusa was recorded from the Antarctic re- gion by Park (1978) as Pseudochirella polyspina. Thirteen species (Table 4) from the Antarctic region have also been collected in the Arctic region (Arctic Ocean basin). They are all bathypelagic, and 77% are rare ex- cept for the following three very common species (num- ber in parenthesis is the number of specimens found in the Southern Ocean by T. Park): Paraeuchaeta barbata (462), Gaetanus tenuispinus (414), and Gaetanus brevispinus (150). The northern and southern polar populations of the species now known as Paraeuchaeta barbata at one time were considered to be a bipolar species, Euchaeta farrani (see Farran, 1929; Vervoort, 1957). Later, Eu- chaeta farrani was synonymized with Euchaeta barbata by Park (1978). Euchaeta barbata was then considered to have a wide distribution throughout the deep water of the world?s oceans, as recorded under that name by Mauchline (1992) and later as Paraeuchaeta barbata by Park (1995). Throughout the Southern Ocean, P. barbata is very common in deep water. Gaetanus tenuispinus is very common south of the Antarctic Convergence. Speci- mens of the third very common species, Gaetanus brevis- pinus Sars, 1900, were initially described from the South- ern Ocean as Gaidius intermedius Wolfenden, 1905, but specimens of this species now are considered to belong to Gaetanus brevispinus (see Markhaseva, 1996). Gaetanus brevispinus is most often encountered in large numbers south of the Antarctic Convergence. 12_Park_pg143-180_Poles.indd 15612_Park_pg143-180_Poles.indd 156 11/17/08 8:30:18 AM11/17/08 8:30:18 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 157 TABLE 5. Abundances and locations of subantarctic deepwater calanoid species absent south of the Antarctic Convergence that occur north of the Subtropical Convergence. 1, south temperate; 2, tropical; 3, north temperate; 4, subarctic; 5, Arctic basin. Abundance in Southern Ocean: CC, very common; C, common; R, rare. A ?H11001? indicates presence. Region Species name Abundance 1 2 3 4 5 Species ranging from subantarctic to south temperate region (6 spp.) Euaugaptilus aliquantus R H11001 Euaugaptilus brevirostratus H11001 Heterorhabdus spinosus CC H11001 Heterorhabdus paraspinosus C H11001 Paraeuchaeta exigua H11001 Scolecithricella pseudopropinqua R H11001 Species ranging from subantarctic to tropical region (2 spp.) Euchirella similis R H11001 H11001 Landrumius gigas H11001 H11001 Species ranging from subantarctic to north temperate region (31 spp.) Aetideus arcuatus H11001 H11001 H11001 Euaugaptilus angustus H11001 H11001 Euaugaptilus gibbus R H11001 H11001 Euaugaptilus laticeps H11001 H11001 H11001 Euaugaptilus oblongus H11001 Euchirella rostrata H11001 H11001 H11001 Gaetanus minor H11001 H11001 H11001 Gaetanus pileatus H11001 H11001 H11001 Heterorhabdus lobatus C H11001 H11001 H11001 Lophothrix frontalis H11001 H11001 H11001 Metridia lucens R H11001 H11001 H11001 Metridia venusta H11001 H11001 H11001 Paraeuchaeta comosa H11001 H11001 H11001 Paraeuchaeta pseudotonsa C H11001 H11001 H11001 Paraeuchaeta sarsi H11001 H11001 H11001 Paraeuchaeta scotti R H11001 H11001 H11001 Pleuromamma abdominalis H11001 H11001 H11001 Pleuromamma peseki H11001 H11001 H11001 Pleuromamma quadrungulata H11001 H11001 H11001 Pleuromamma xiphias H11001 H11001 H11001 Scaphocalanus cristatus R H11001 Scaphocalanus echinatus CC H11001 H11001 H11001 Scaphocalanus medius C H11001 H11001 Scolecithricella dentata H11001 H11001 H11001 Scolecithricella profunda H11001 H11001 H11001 Scolecithricella vittata R H11001 H11001 H11001 Scottocalanus securifrons H11001 H11001 H11001 Scottocalanus helenae H11001 H11001 H11001 Undeuchaeta major H11001 H11001 H11001 Undeuchaeta plumosa H11001 H11001 H11001 Valdiviella minor H11001 H11001 H11001 Species ranging from subantarctic to subarctic waters (10 spp.) Centraugaptilus rattrayi R H11001 H11001 H11001 H11001 Chirundina streetsii H11001 H11001 H11001 H11001 Disseta palumbii H11001 H11001 H11001 H11001 Euchirella maxima H11001 H11001 H11001 H11001 Gaetanus kruppii H11001 H11001 H11001 H11001 Gaetanus latifrons R H11001 H11001 H11001 H11001 Metridia brevicauda H11001 H11001 H11001 H11001 Paraeuchaeta hansenii H11001 H11001 H11001 H11001 Scottocalanus thorii H11001 H11001 Undinella brevipes H11001 H11001 Species ranging from subantarctic to Arctic basin (1 sp.) Paraheterorhabdus compactus R H11001 H11001 H11001 H11001 12_Park_pg143-180_Poles.indd 15712_Park_pg143-180_Poles.indd 157 11/17/08 8:30:19 AM11/17/08 8:30:19 AM 158 SMITHSONIAN AT THE POLES / PARK AND FERRARI In summary, there are 84 species of deepwater calanoid copepods that occur south of the Antarctic Convergence and have also been reported northward to different degrees; 86% of these species have been reported at least as far north as the north temperate region. Two of the seven species oc- curring from the Antarctic region to the south temperate re- gion are common or very common in the Southern Ocean, while one of the fi ve species found from the Antarctic region north to the tropical region is very common in the Southern Ocean. Seven of the 29 species reported from the Antarc- tic region and the north temperate region are common or very common in the Southern Ocean. Four of the 30 spe- cies from the Antarctic region and reported as far north as the subarctic region are common or very common in the Southern Ocean. Only 3 of the 13 species found in the Arc- tic Ocean are common or very common in the Southern Ocean. These observations suggest that most (80%) of the rare or very rare deepwater species occurring in the Antarc- tic region appear to be distributed widely throughout the world?s oceans, where they also are rare or very rare deep- water species. However, there are a small number (17) of deepwater species that may be collected in large numbers in the Southern Ocean that are widely distributed and rare or very rare outside of the Southern Ocean. In contrast to the epipelagic calanoid community, the deepwater pelagic calanoid community of the Southern Ocean is represented by a very diverse group of species. Many of the endemic species collected in the Southern Ocean are common or very common there, apparently having adapted to the high primary and secondary pro- ductivity (Park, 1994). Interestingly, a few of the species collected in other regions are also very common in the Southern Ocean, although they are known from only a few specimens throughout the rest of their range. These species appear to be capable of surviving in habitats of low productivity, and yet they can maintain larger popula- tions in some eutrophic habitats like the Southern Ocean. This small number of deepwater species of pelagic cala- noid copepods may also be well adapted to high primary and secondary productivity of the Southern Ocean, and this adaptation may result in relatively larger numbers of specimens (Park, 1994). SUBANTARCTIC DEEPWATER CALANOIDS REPORTED NORTH OF THE SUBTROPICAL CONVERGENCE Of the 167 deepwater calanoid species found in the subantarctic region, between the Antarctic Convergence and Subtropical Convergence, seven are endemic to that region; 110 species were also collected south of the Ant- arctic Convergence. The remaining 50 species are not col- lected south of the Antarctic Convergence, but their distri- bution does extend north of the Subtropical Convergence (Table 5). Six species (12%) can be found in the south temperate region, and two (4%) have been collected in the tropical region. There are records of 31 species (62%) in the north temperate region, 10 species (20%) in the sub- arctic region, and 1 species (2%) from the Arctic basin. Of the six species ranging from the subantarctic to the south temperate regions, Heterorhabdus spinosus is very common, and H. paraspinosus and Paraeuchaeta exigua are common in the subantarctic region. The two species of Heterorhabdus occur together and have been collected from only three other widely separated areas: off the west coast of South Africa, off the southern west coast of Chile, and off the east coast of New Zealand and in the Tasman Sea. Paraeuchaeta exigua has been found in four widely separated areas: the eastern and the western parts of the South Atlantic Ocean, the Tasman Sea, and the western Indian Ocean, where it is very com- mon (Park, 1995). These three species apparently are as- sociated with habitats of high secondary productivity, es- pecially coastal upwelling systems. The remaining three species have been found only once and remain poorly known. Only two species, Euchirella similis and Landrumius gigas, have been reported from the subantarctic region to the tropical region. They are very rare in the subantarctic region and remain poorly known. All 31 species rang- ing from the subantarctic region to the north temperate region were originally described from the low or middle latitudes and subsequently have been reported from the subantarctic region. All are believed to be mesopelagic or bathypelagic except for two relatively shallow living species, Scolecithricella dentata and Scolecithricella vit- tata. Twenty-fi ve of the species are rare in the subant- arctic region, and fi ve species, Scottocalanus securifrons, Paraeuchaeta pseudotonsa, Lophothrix frontalis, Scaph- ocalanus medius, and Heterorhabdus lobatus, are com- mon. Only Scaphocalanus echinatus is very common in the subantarctic region. Ten of the subantarctic species have been found as far north as the subarctic region. They are all rare deepwater calanoids. Paraheterorhabdus compactus is the only spe- cies known to occur from the subantarctic region north to the Arctic basin; it is bathypelagic and occurs in small numbers throughout its range (Park, 2000). In summary, there are 50 species that are absent south of the Antarctic Convergence but are found in the subant- arctic region and northward to varying degrees. Eighty- 12_Park_pg143-180_Poles.indd 15812_Park_pg143-180_Poles.indd 158 11/17/08 8:30:19 AM11/17/08 8:30:19 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 159 four percent of these species have been reported to at least the north temperate region. Eighty-two percent are rare; the exceptions are three of six species occurring in the subantarctic and south temperate regions, where they are common in the productive coastal waters, and 6 of 31 spe- cies found from the subantarctic to the north temperate regions. These latter six are common or very common in productive the subantarctic region. SOUTHERN OCEAN CALANOIDS WITH A BIPOLAR DISTRIBUTION There are nine pelagic calanoids whose distribution can be described as bipolar (Table 6). Aetideopsis minor, Pseudochirella spectabilis, and Spinocalanus antarcti- cus are found south of the Antarctic Convergence and in the Arctic basin (Table 4); Aetideopsis rostrata and Pseudochirella batillipa are found south of the Antarctic Convergence, in the Arctic basin and its adjacent boreal seas (Table 4); Metridia ornata and Racovitzanus ant- arcticus are found south of the Antarctic Convergence and in boreal seas adjacent to the Arctic basin (Table 4); Batheuchaeta peculiaris and Chiridius polaris are found both north and south of the Antarctic Convergence and in boreal seas adjacent to the Arctic basin. Eight of these nine species are rare or very rare deepwater spe- cies in both polar areas, and three of those eight have been found in the subarctic region but not in the Arctic Ocean basin. The ninth species, Racovitzanus antarcti- cus, is very common in the waters south of the Antarctic Convergence and has been described as common in the boreal seas adjacent to the Arctic basin (Brodsky, 1950). Park (1983a) examined specimens of R. antarcticus from the northern Pacifi c Ocean and found that they are iden- tical to those from the Southern Ocean in anatomical details of the exoskeleton. In the Southern Ocean, the number of R. antarcticus collected appears to decrease rather abruptly with distance northward from the Ant- arctic Convergence, and there is some evidence that the species may be found in deeper waters north of the con- vergence but within the Southern Ocean (Park, 1983a). In the Northern Hemisphere, R. antarcticus seems to in- habit the mesopelagic zone (200? 1,000 m). Two hypotheses can be suggested to explain the dis- tribution of R. antarcticus. The polar populations may be connected through very deep living populations in the north temperate, tropical, and south temperate regions at depths not adequately sampled to date. This connection would mediate gene fl ow through the undetected deepwa- ter populations in the temperate and tropical regions and would result in the stable morphological similarity between specimens from the Southern and Northern hemispheres. A similar scenario may explain the apparent bipolar dis- tribution of remaining eight deepwater calanoid copepods that are rare or very rare: at high latitudes they inhabit shallower depths, where individuals can be captured more easily, perhaps because secondary productivity is higher. At lower latitudes, populations are found much deeper and are not as readily collected. The alternate hypothesis is of incipient speciation from a previously more broadly distributed deepwater species that is no longer connected through temperate and tropical deepwater populations (see the Comparative Endemicity of the Southern Ocean Fauna section). Morphological similarity in this case is transitory because the absence of gene fl ow between the polar populations is expected to result in morphological divergence. The remaining eight rare species have come to be rec- ognized as having a bipolar distribution in one of three ways. Aetideopsis minor, Chiridius polaris, Pseudochire- lla batillipa, and Spinocalanus antarcticus originally were described from the Southern Ocean and subsequently reported from the Northern Hemisphere. Spinocalanus antarcticus was discovered in the Arctic Ocean ( Damkaer, 1975), while Aetideopsis minor, Chiridius polaris, and Pseudochirella batillipa recently have been recorded for the fi rst time beyond their type locality, in the Arc- tic Ocean and adjacent boreal seas (Markhaseva, 1996). Two species, Batheuchaeta peculiaris and Metridia ornata, originally were described from localities adjacent to the Arctic Ocean; subsequently, they were reported from the Southern Ocean, for the fi rst time outside their type local- ity, by Markhaseva (1996, 2001). Finally, two species were TABLE 6. Nine Calanoid species with a bipolar distribution. Species name Distribution Aetideopsis minor Antarctic (61?? 69?S), Arctic basin Aetideopsis rostrata Antarctic, Arctic and boreal seas Batheuchaeta peculiaris Antarctic (63?S), boreal region (45?? 46?N) Chiridius polaris Antarctic (53?? 68?S), boreal region (44?? 46?N) Metridia ornata Antarctic (55?? 70?S), boreal region (38?? 57?N) Pseudochirella batillipa Antarctic (53?? 66?S), 86?N and 44?? 46?N Pseudochirella spectabilis Antarctic (61?? 68?S), Arctic basin Racovitzanus antarcticus Southern Ocean, boreal seas Spinocalanus antarcticus Antarctic, Arctic basin 12_Park_pg143-180_Poles.indd 15912_Park_pg143-180_Poles.indd 159 11/17/08 8:30:20 AM11/17/08 8:30:20 AM 160 SMITHSONIAN AT THE POLES / PARK AND FERRARI recognized to be bipolar when specimens from southern and northern localities, originally considered different spe- cies, subsequently were proposed to be identical. Aetideop- sis infl ata, originally described from the Antarctic region, was synonymized with the subarctic species Aetideopsis rostrata (see Markhaseva, 1996), so Aetideopsis rostrata is now a bipolar species. Similarly, Pseudochirella elon- gata, also originally described from the Antarctic region, was synonymized with the Arctic species Pseudochirella spectabilis (see Markhaseva, 1996); the latter species now has a bipolar distribution. The geographical distribution of these rare species, as inferred from a small number of specimens and from a limited number of localities, how- ever, may not be completely understood. It is worth noting that some species of pelagic calanoid copepods previously regarded as having disjunct popula- tions in the Southern and Arctic oceans have not subse- quently been found to be bipolar. Rather, the northern and southern populations have been recognized as two sepa- rate species. As examples, the southern population pre- viously referred to as Calanus fi nmarchicus is now Cala- nus australis; the northern population previously known as Neocalanus tonsus is now Neocalanus plumchrus; the southern population originally known as Scaphocalanus brevicornis is now Scaphocalanus farrani. The fi rst two species are epipelagic herbivores; the third is a deepwa- ter detritivore. The taxonomic history of Paraeuchaeta barbata is informative but more complex (see the Deep- water Calanoids from Antarctic Waters Reported North of the Antarctic Convergence section). This species origi- nally was described as Euchaeta farrani from the Norwe- gian Sea by With (1915); subsequently, it was recorded from the Antarctic region by Farran (1929) and Vervoort (1957) and proposed by them to be a species with a bipo- lar distribution. As described more completely above (see the Deepwater Calanoids from Antarctic Waters Reported North of the Antarctic Convergence section), these speci- mens have been recognized by Park (1995) as belonging to P. barbata, a deepwater carnivore, now understood to be distributed throughout the world?s oceans. In summary, taxonomic analyses have reversed initial inferences of a bipolar distribution for Calanus fi nmarchi- cus, Neocalanus tonsus, and Scaphocalanus brevicornis. However, taxonomic analyses have established a bipolar distribution for the rare and very rare deepwater species Aetideopsis rostrata, Aetideopsis minor, Chiridius polaris, Pseudochirella spectabilis, and Pseudochirella batillipa. The distribution of the very common P. barbata offers rea- sons for caution in hypothesizing a bipolar distribution for rare and very rare deepwater species. VERY COMMON PELAGIC CALANOIDS AND AREAS OF HIGH PRODUCTIVITY All of the very common epipelagic calanoids of the Southern Ocean are herbivores (Table 2). In order of abundance, they are Calanoides acutus, Rhincalanus gi- gas, Calanus propinquus, Calanus simillimus, Metridia gerlachei, Clausocalanus laticeps, and Clausocalanus brevipes. These epipelagic calanoids are endemic to the Southern Ocean and appear to have successfully adapted to the high primary productivity there. The eutrophic con- ditions there may also be responsible for the high numbers of individuals of these endemic herbivores. The seven epi- pelagic species together make up the enormous secondary biomass of the Southern Ocean, a secondary biomass that is unsurpassed in any other region of the world?s oceans (Foxton, 1956). Among the deepwater calanoids of the Southern Ocean, there are 26 species (Table 7) from the studies of Park (1978, 1980, 1982, 1983a, 1983b, 1988, 1993) that are represented by more than 100 individuals and are considered very common. A majority, 14 of 26 species, of these very common deepwater calanoids are limited in their distribution to the Southern Ocean. Among the 14 very common deepwater endemic species, eight belong to two genera in the family Scolecitrichidae (fi ve species of Scaphocalanus and three of Scolecithricella); all are de- tritivores. These are followed, in order of abundance, by three species of Paraeuchaeta in the Euchaetidae. Twelve very common species are more widely distrib- uted, found northward at varying distances beyond the Subtropical Convergence. Four species have been found as far north as the north temperate region, one has been found in the subarctic region, and three other deepwater species have a range extending into the Arctic Ocean. Ra- covitzanus antarcticus has a bipolar distribution. Most of these very common species, then, are either endemic to the Southern Ocean (14 species) or have a broad distribution extending north of the tropical region (9 species). Among the 26 common deepwater calanoids of the Southern Ocean, 12 species belong to the family Scolecit- richidae (six species of Scaphocalanus, fi ve species of Scole- cithricella, and one species of Racovitzanus). The family is followed, in order of number of species, by the families Eu- chaetidae, with fi ve species all belonging to the genus Par- aeuchaeta; Aetideidae, with three species (two of Gaetanus and one Euchirella); Augaptilidae, also with three species (two Haloptilus and one Euaugaptilus); Phaennidae, with two species (both Cornucalanus); and Heterorhabdidae, with one species belonging to the genus Heterorhabdus. 12_Park_pg143-180_Poles.indd 16012_Park_pg143-180_Poles.indd 160 11/17/08 8:30:20 AM11/17/08 8:30:20 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 161 The 12 scolecitrichid species together were represented by 9,642 individuals from the Southern Ocean, and the fi ve Paraeuchaeta species were represented by 2,089 individu- als. Three aetideid species and three augaptilids were repre- sented by 746 and 676 individuals, respectively. On the basis of the relatively large number of speci- mens of Paraeuchaeta, Aetideidae, Heterorhabdidae, and Augaptilidae that are carnivores, all of these species are presumed to be well adapted to the high secondary pro- ductivity resulting from the large populations of epipelagic herbivores in the Southern Ocean. Species of Scolecitrichi- dae play a major ecological role as pelagic detritivores in the Southern Ocean, just as species of Scolecitrichidae and related bradfordian families of calanoids play a similar role (detritivory) in the deepwater benthopelagic habitat of other oceans (Markhaseva and Ferrari, 2006). Because of their relatively small body size, scolecitrichids may also serve as a food source for carnivorous calanoids like spe- cies of Paraeuchaeta, the aetideids, and the augaptilids during periods when the juvenile stages of herbivores are unavailable as prey for these carnivores. These conclusions are reinforced by restricting ob- servations to the 10 species represented in the Southern Ocean by more than 400 individuals (with number of indi- viduals in parenthesis): Scaphocalanus vervoorti (1,936), Scolecithricella minor (1,728), S. dentipes (1,603), Scaphocalanus farrani (1,271), Racovitzanus antarcticus (1,077), Scolecithricella cenotelis (929), Paraeuchaeta antarctica (602), Paraeuchaeta rasa (546), Paraeuchaeta barbata (462), and Gaetanus tenuispinus (414). Four of the scolecitrichid species, Scaphocalanus vervoorti, S. far- rani, Scolecithricella dentipes, and S. cenotelis, and two of the euchaetid species, Paraeuchaeta antarctica and P. rasa, are endemic to the Southern Ocean. The range of the very common Scolecithricella minor extends into the sub- arctic region, while Paraeuchaeta barbata and Gaetanus tenuispinus have been collected as far north as the Arctic basin. The scolecitrichid Racovitzanus antarcticus is also among the most common species in the Southern Ocean but exhibits a bipolar distribution, occurring in boreal wa- ters adjacent to the Arctic basin. These 10 very common species either are endemic to the Southern Ocean (Scapho- calanus vervoorti, Scolecithricella dentipes, Scaphocalanus farrani, Scolecithricella cenotelis, Paraeuchaeta antarctica, and Paraeuchaeta rasa) or have a distribution that extends as far north as the subarctic region or Arctic basin (Scole- cithricella minor, Racovitzanus antarctica, Paraeuchaeta barbata, and Gaetanus tenuispinus). None of the very common deepwater Southern Ocean pelagic calanoids have distributions that extend only to the south temperate region to the tropical region. Within the Southern Ocean the abundance and dis- tribution of deepwater calanoids are believed to be de- termined, for the most part, by the availability of food rather than their adaptation to nonbiological environ- mental parameters such as water temperature (Park, 1994). Whether these eutrophic species are endemics or not, they are restricted to water of high primary and sec- ondary productivity. They can be expected to be common or very common due to their adaptations for exploiting TABLE 7. Very common deepwater calanoid species of the Southern Ocean (26 spp.). Species name Number of of specimens Species endemic to the Southern Ocean (14 spp.) Euaugaptilus antarcticus 136 Euchirella rostromagna 182 Haloptilus ocellatus 152 Paraeuchaeta antarctica 602 Paraeuchaeta biloba 370 Paraeuchaeta rasa 546 Scaphocalanus antarcticus 130 Scaphocalanus farrani 1,271 Scaphocalanus parantarcticus 289 Scaphocalanus subbrevicornis 188 Scaphocalanus vervoorti 1,936 Scolecithricella cenotelis 929 Scolecithricella dentipes 1,603 Scolecithricella schizosoma 151 Species ranging from the Southern Ocean to south temperate region (2 spp.) Paraeuchaeta regalis 109 Heterorhabdus spinosus 243 Species ranging from the Southern Ocean to the tropical region (1 sp.) Cornucalanus robustus 161 Species ranging from the Southern Ocean to the north temperate region (4 spp.) Haloptilus oxycephalus 388 Scolecithricella emarginata 226 Cornucalanus chelifer 111 Scaphocalanus echinatus 114 Species ranging from the Southern Ocean to the subarctic region (1 sp.) Scolecithricella minor 1,728 Species ranging from the Southern Ocean to the Arctic basin (3 spp.) Gaetanus brevispinus 150 Gaetanus tenuispinus 414 Paraeuchaeta barbata 462 Species with a bipolar distribution (1 sp.) Racovitzanus antarcticus 1,077 12_Park_pg143-180_Poles.indd 16112_Park_pg143-180_Poles.indd 161 11/17/08 8:30:21 AM11/17/08 8:30:21 AM 162 SMITHSONIAN AT THE POLES / PARK AND FERRARI the available food sources associated with that habitat. In contrast, oligotrophic species in the Southern Ocean are not presumed to be adapted to waters of high pro- ductivity. Their distribution is expected to be worldwide because they are capable of surviving at most levels of food resources anywhere in the world?s oceans. How- ever, oligotrophic species are expected to be rare or very rare in most regions of the world?s oceans. With these constraints, a distribution of common or very common species is expected to be limited to the highly produc- tive Southern Ocean; this is observed about half the time. Fourteen of the 26 common or very common species of the Southern Ocean are endemic. As noted, among the 10 most numerous of the very common species found in the Southern Ocean, six are en- demic. Of the remaining four species represented by more than 400 specimens in the Southern Ocean, Scolecithri- cella minor, Gaetanus tenuispinus, and Paraeuchaeta bar- bata have been reported throughout the world?s oceans, while Racovitzanus antarctica appears to have a bipolar distribution, restricted to the Southern Ocean and to the subarctic region (boreal seas adjacent to the Arctic Ocean). The distribution of Scolecithricella minor and Gaetanus tenuispinus outside the Southern Ocean is based on litera- ture records. Until these records can be verifi ed by direct comparison of specimens, the relationship of the South- ern Ocean specimens to specimens collected elsewhere re- mains tentative, and we are unable to contribute more to the nature of these distributions. The distribution of Paraeuchaeta barbata has become clearer in recent years and can also be understood within the context of the association of this abundant species with areas of high primary and secondary productivity. The polar populations of Paraeuchaeta barbata were once regarded as a separate, bipolar species (see the Deepwa- ter Calanoids from Antarctic Waters Reported north of the Antarctic Convergence section). When Park (1995) restudied the various populations by analyzing a large number of specimens collected throughout the world?s oceans, he found that specimens exhibited a considerable but continuous variation in size. As a result of this analysis and an earlier restricted analysis (Mauchline, 1992), body size was rejected as species-specifi c character state for P. barbata. This considerable and continuous variation in body size of P. barbata was subsequently reexamined in asso- ciation with the distribution of this species (T. Park, un- published observations). Large-sized individuals occur not only at the high latitudes of both hemispheres but also along the west coast of the Americas in areas associated with signifi cant coastal upwelling systems. Large individu- als also were recorded in the Malay Archipelago and along the east coast of Japan up to Kuril and Kamchatka; these are also seasonally episodic areas of upwelling. Coastal upwelling systems along the west coast of the Americas and the east coast of Japan result in high primary and secondary productivity, which, in turn, may explain the larger-size individuals of P. barbata in these areas. The smallest individuals of P. barbata are found in the middle of the North Atlantic, an oligotrophic habitat. Species like P. barbata, which are distributed through- out the world?s oceans, may have become very common in the Southern Ocean and other areas of seasonally epi- sodic upwelling by taking advantage of the high second- ary productivity of eutrophic habitats; individuals of this species are also larger in these habitats as a result of the availability of prey. In contrast, away from areas of high productivity, few specimens are collected, and individuals are smaller in size. COMPARATIVE ENDEMICITY OF THE SOUTHERN OCEAN FAUNA The pelagic calanoid families Euchaetidae and Het- erorhabdidae have been studied throughout the world?s oceans (Park, 1995, 2000). From these publications, the number of endemic species belonging to these two families from the Southern Ocean can be compared to the num- ber of endemics from three other areas of interest of the world?s oceans: Arctic-boreal (including the adjacent bo- real seas of the Atlantic and Pacifi c Oceans), Indo-West Pacifi c, and eastern Pacifi c. The Southern Ocean, with 10 endemic species of Paraeuchaeta, has the highest number for that genus (Table 8), followed by the Arctic Ocean, with seven endemic species of Paraeuchaeta, and the Indo- West Pacifi c and the eastern Pacifi c, each with four en- demic species. Twenty-three of the 25 endemic species of Paraeuchaeta referred to above are bathypelagic; the ex- ceptions are the Indo-West Pacifi c epipelagic species Par- aeuchaeta russelli and P. simplex. Six of the 10 Southern Ocean endemics, P. antarctica, P. biloba, P. dactylifera, P. eltaninae, P. parvula, and P. rasa, have been found in large numbers. Paraeuchaeta aus- trina, P. erebi, and P. tycodesma have not been collected in large numbers, but they appear to be restricted to the ice edge along Antarctica. This habitat may not have been adequately sampled with midwater trawls, and as a result, these species may be underrepresented in trawl samples. Paraeuchaeta similis and P. antarctica have a broader dis- 12_Park_pg143-180_Poles.indd 16212_Park_pg143-180_Poles.indd 162 11/17/08 8:30:21 AM11/17/08 8:30:21 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 163 TABLE 8. Endemic species of Paraeuchaeta and Heterorhabdidae found in four different areas of interest. A ?H11001? indicates presence. Area of interest Species Southern Ocean Arctic-boreal Eastern Pacifi c Indo-West Pacifi c Paraeuchaeta antarctica H11001 P. austrina H11001 P. biloba H11001 P. dactylifera H11001 P. eltaninae H11001 P. erebi H11001 P. parvula H11001 P. rasa H11001 P. similis H11001 P. tycodesma H11001 P. birostrata H11001 P. brevirostris H11001 P. elongata H11001 P. glacialis H11001 P. norvegica H11001 P. polaris H11001 P. rubra H11001 P. californica H11001 P. copleyae H11001 P. grandiremis H11001 P. papilliger H11001 P. eminens H11001 P. investigatoris H11001 P. russelli H11001 P. simplex H11001 Heterorhabdus austrinus H11001 H. pustulifer H11001 H. spinosus H11001 H. paraspinosus H11001 Heterostylites nigrotinctus H11001 Paraheterorhabdus farrani H11001 Heterorhabdus fi stulosus H11001 H. norvegicus H11001 H. tanneri H11001 Paraheterorhabdus longispinus H11001 Heterorhabdus abyssalis H11001 H. americanus H11001 H. prolixus H11001 H. quadrilobus H11001 Heterostylites echinatus H11001 tribution within the Southern Ocean, but only P. antarc- tica has been collected in large numbers. The endemic species of the Arctic Ocean, including adjacent boreal waters, and the endemics of the eastern Pacifi c have also been found in large numbers. These spe- cies are all believed to inhabit waters of high primary and secondary productivity, where endemism may have devel- oped as an adaptation to these eutrophic habitats (Park, 1994). Of the four endemics of the Indo-West Pacifi c, Paraeuchaeta russelli and P. simplex are neritic, inhabit- ing relatively shallow water. Paraeuchaeta eminens and P. investigatoris are deepwater species. All four species are common in waters of the Malay Archipelago, an area with relatively high primary and secondary productivity. High primary and secondary productivity, rather than a habi- tats abiological attributes, appears to have been the pri- mary determinant for the evolution of endemicity among these species of Paraeuchaeta. Within the family Heterorhabdidae, six species are en- demic to the Southern Ocean as compared to fi ve endemic 12_Park_pg143-180_Poles.indd 16312_Park_pg143-180_Poles.indd 163 11/17/08 8:30:22 AM11/17/08 8:30:22 AM 164 SMITHSONIAN AT THE POLES / PARK AND FERRARI species in the eastern Pacifi c. Four species are restricted to the Arctic-boreal area, including three species endemic to the boreal Pacifi c and one endemic species found in the boreal Atlantic. No endemic species of Heterorhabdidae is found in the Indo-West Pacifi c. All of the heterorhabdid species discussed here are assumed to be carnivores, with the exception of Heterostylites echinatus (see Ohtsuka et al., 1997), and carnivory is assumed to have arisen from suspension feeding within the Heterorhabdidae only once (Ohtsuka et al., 1997). The highest number of endemic heterorhabdid species, like the number of endemics of Paraeuchaeta, is found in the Southern Ocean. However, in contrast to species of Par- aeuchaeta, of the six Southern Ocean endemic heterorhab- dids, only Heterorhabdus spinosus is found in large num- bers; it is common in coastal waters. Beyond the Southern Ocean, among the four endemic species of the Arctic and boreal seas, Heterorhabdus norvegicus is very common in the boreal Atlantic. Heterorhabdus fi stulosus, H. tanneri, and Paraheterorhabdus longispinus are very common along the coasts of the boreal Pacifi c, and all occur in large num- bers in some localities. Among the fi ve endemic species of Heterorhabdidae in the eastern Pacifi c, all are limited in their distribution to waters close to the coasts of Ameri- cas, in areas of coastal upwelling, where they have been found in large numbers. One explanation for the different occurrences of Paraeuchaeta and heterorhabdid endemics in the Southern Ocean is that the heterorhabdids may be relatively late colonizers of the Southern Ocean; species of Paraeuchaeta may already have established themselves as the dominant carnivores before colonization of the South- ern Ocean by species of Heterorhabdidae. In summary, within the two families of pelagic cala- noids that have been studied worldwide, the highest num- ber of endemic species is found in the Southern Ocean. All Southern Ocean endemics of Paraeuchaeta are found in large numbers, except for P. similis and three species found near the ice edge adjacent to Antarctica where these ice edge species may have been undersampled. In contrast, the six endemic species of Heterorhabdidae found in the Southern Ocean are rare. However, beyond the Southern Ocean the endemics of both families of carnivores are as- sociated with the waters of high primary and secondary productivity, where they may be collected in large num- bers. On the basis of these observations, the endemicity of many of the very common bathypelagic calanoids of the Southern Ocean, like the endemicity of Southern Ocean epipelagic calanoids, is suggested to have resulted from the adaptation to conditions of high primary and secondary productivity. EVOLUTION OF THE PELAGIC CALANOID FAUNA WITHIN THE SOUTHERN OCEAN Among the 184 deepwater calanoid species found in the Southern Ocean, 50 species (27%) occur exclusively in the Southern Ocean; 20 of those species (40%) are rare or very rare (Table 3). Several factors may be responsible for the evolution of the deepwater pelagic calanoid fauna, re- straining their dispersal throughout the deepwater of the world?s oceans and selecting for this endemicity. Water tem- perature, as represented by the rather abrupt changes at the Antarctic Convergence or the Subtropical Convergence, is unlikely to affect the structure or the distribution of the deepwater calanoids because water temperatures below 1,000 m are uniformly cold within the Southern Ocean, and this uniformly cold deepwater is continuous with the deep- water of the adjacent Pacifi c, Atlantic, and Indian oceans. The proposed relationship between habitat productivity and endemism may be a more useful initial condition. The majority (60%) of deepwater endemic species of the South- ern Ocean are common or very common. As mentioned earlier, this may be the product of the high primary and secondary productivity of the Southern Ocean, especially south of the Antarctic Convergence, resulting in the evolu- tion and adaptation of an oligotrophic species to this eu- trophic habitat. Endemism of deepwater pelagic calanoids in the Southern Ocean, therefore, is hypothesized to have evolved as rare species that are widely distributed in oli- gotrophic habitats throughout the world?s oceans became adapted to exploit high primary and secondary productive habitats (Park, 1994); these adaptations have resulted in an increased population size of the eutrophic species. A second explanatory condition for the evolution of the pelagic, marine calanoid fauna in the Southern Ocean depends on whether polar species within a single genus are monophyletic, having evolved from a single ancestral spe- cies that initially colonized the Southern Ocean, or polyphy- letic, with each species having evolved independently from an ancestor distributed outside of the Southern Ocean or by evolving from more than one initially colonized ancestral species. There is evidence that supports this latter model of independent colonizations for Southern Ocean endemic species of the families Euchaetidae and Heterorhabdidae (see the Comparative Endemicity of the Southern Ocean Fauna section), although the situation is more complex for the antarctica species group of Paraeuchaeta. Further evidence can be found in the phenomenon of sibling species pairs. When morphological details are closely compared, one species often can be found outside the Southern Ocean that is very similar to each Southern 12_Park_pg143-180_Poles.indd 16412_Park_pg143-180_Poles.indd 164 11/17/08 8:30:22 AM11/17/08 8:30:22 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 165 Ocean endemic. These two species, the Southern Ocean endemic and its closest relative outside of the Southern Ocean, are referred to here as a sibling species pair. Fifteen endemics among the 17 sibling species pairs (Figure 1) have an allopatric distribution, rather than being sympat- ric with its closest relative. In addition, the antarctica spe- cies group of Paraeuchaeta is also allopatric with its most closely related congener, P. bisinuata. The only exception is the pair Haloptilus ocellatus and H. oxycephalus; these two species may be considered allopatric but with a narrow Calanoides acutus C. carinaatus Euchirella rostromagna E. rostrata Haloptilus ocellatus H. oxycephalus Heterorhabdus austrinus H. abyssalis Onchocalanus paratrigoniceps O. trigoniceps Paraeuchaeta rasa P. aequatorialis P. antarctica sp gr P. bisinuata Paraeuchaeta regalis P. calva Paraeuchaeta eltaninae P. scotti Paraeuchaeta pseudotonsa P. tonsa Paraheterorhabdus farrani P. robustus Pseudochirella mawsoni P. pustulifera Rhincalanus gigas R. nasutus Scaphocalanus antarcticus S. cristatus Scaphocalanus farrani S. echinatus Scolecithricella hadrosoma S. emarginata Scolecithricella cenotelis S. ovata S 35 0 35 N FIGURE 1. Distribution of selected pelagic calanoids of the Southern Ocean and their closest relatives. 12_Park_pg143-180_Poles.indd 16512_Park_pg143-180_Poles.indd 165 11/17/08 8:30:23 AM11/17/08 8:30:23 AM 166 SMITHSONIAN AT THE POLES / PARK AND FERRARI zone of overlap. For many common or very common deep- water calanoid species that are endemic to the Southern Ocean (Figure 1, Tables 4, 5), the closest relative is not found in the Southern Ocean, is widely distributed, is rare, and is usually associated with oligotrophic habitats. Ex- amples of these deepwater sibling species pairs (Southern Ocean species fi rst) include (Figure 1) Euchirella rostro- magna and E. rostrata, Paraeuchaeta pseudotonsa and P. tonsa, P. antarctica and P. bisinuata, Scaphocalanus crista- tus and S. antarcticus, and Paraheterorhabdus farrani and P. robustus. The members of some sibling species pairs are so similar to each other morphologically that, originally, they were regarded as the same species, e.g., Paraeuchaeta tonsa and P. pseudotonsa or Euchirella rostrata and E. rostromagna. In addition to being more numerous, the Southern Ocean endemics are usually larger in size than their smaller, rare, cosmopolitan counterparts. Between members of the deepwater pairs, the common or very common endemics adapted to the eutrophic habitat of the Southern Ocean are hypothesized to have evolved from a rare widespread species adapted to oligotrophic habi- tats (Park, 1994). This evolutionary event has resulted in two closely related species, a sibling species pair with the species outside of the Southern Ocean remaining adapted to an oligotrophic environment and the Southern Ocean species adapted to high-productivity habitats. In view of the close morphological similarity between the species of a sibling species pair, this process seems to have a relatively short evolutionary history. However, this hypothesis does not imply that all pairs evolved about the same time. The situation for Paraeuchaeta antarctica appears to be more complicated than a case of a simple sibling species pair. Paraeuchaeta antarctica is a very common predator and is morphologically similar to four other endemic spe- cies, P. similis, P. austrina, P. erebi, and P. tycodesma. These fi ve endemic species make up the antarctica species group (Fontaine, 1988). All fi ve of these species can be found to occur sympatrically adjacent to the ice edge of Antarctica. Paraeuchaeta austrina, P. erebi, and P. tycodesma are re- stricted to this habitat, while P. antarctica and P. similis may be found throughout the Southern Ocean. The most similar congener, and presumed closest relative, of the antarctica species group is P. bisinuata. Paraeuchaeta bi- sinuata is a rare deepwater species found in all the world?s oceans except the Southern Ocean. Paraeuchaeta bisinu- ata and the common ancestor of the antarctica species group are hypothesized to have been a sibling species pair. The common ancestor of the antarctica species group is assumed to have colonized the Southern Ocean, eventu- ally adapting and being confi ned to the eutrophic habitat. All of its descendants, including P. austrina, P. erebi, and P. tycodesma, which are associated with waters adjacent to the ice edge, and the more broadly distributed P. similis and P. antarctica, are restricted to the Southern Ocean. Paraeuchaeta bisinuata, the cosmopolitan species of the original pair, remains associated with oligotrophic habi- tats throughout the world?s oceans. To summarize, the evolution of deepwater endemic species of the Southern Ocean can be hypothesized from an ordered set of changes in distribution and subsequent morphological divergence in the following way: (1) be- ginning with a rare, widely distributed species adapted to oligotrophic habitats, e.g., Augaptilus glacialis, (2) a Southern Ocean population becomes associated with its eutrophic habitat and becomes separated from the rare, widely distributed, oligotrophic species; (3) the Southern Ocean endemic population adapts to this eutrophic habi- tat, and its population size increases. It diverges from the rare, widely distributed, oligotrophic species, resulting in a sibling species pair, e.g., Scolecithricella farrani and S. echinates. Another type of species pair identifi ed in this study requires different explanatory conditions about the evolu- tion of the Southern Ocean fauna. A bipolar species pair consists of two morphologically similar species, presumed closest relatives, one that is endemic to the Southern Ocean and a second that is endemic to the Arctic Ocean and adjacent boreal waters. Several endemic species of the Southern Ocean have a morphologically similar congener in the Arctic region (Table 7, Figure 2). The morphological similarities between the members are so close that some of the pairs were recognized as separate species only recently, e.g., the Southern Ocean Scaphocalanus farrani was sepa- rated from S. brevicornis by Park (1982). In general, the number of morphological differences is few and the degree of the morphological divergence is slight between members of these southern and northern oceanic pairs, e.g., Para- heterorhabdus farrani and P. longispinus, Scaphocalanus parantarcticus and S. acrocephalus, and Paraeuchaeta re- galis and P. polaris. The extent of the morphological simi- larity between these polar species suggests that they may have been derived from a common ancestor (see below), although this does not imply that all pairs have evolved about the same time. The evolution of deepwater bipolar species pairs can be hypothesized from an ordered set of changes in dis- tribution and subsequent morphological divergence in the following way. Beginning with a widely distributed deep- water species with shallow populations in the Southern Ocean and Arctic Ocean, e.g., Paraeuchaeta barbata, (1) 12_Park_pg143-180_Poles.indd 16612_Park_pg143-180_Poles.indd 166 11/17/08 8:30:24 AM11/17/08 8:30:24 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 167 deepwater middle- and low-latitude populations become extinct over varying periods of time, resulting in a non- uniform distribution, e.g., Batheuchaeta peculiaris, (2) complete middle- and low-latitude extinctions eventually result in a species with a bipolar distribution, e.g., Pseu- dochirella spectabilis, and (3) subsequent morphological divergence of the bipolar populations results in a bipolar pair of species, e.g., Paraeuchaeta regalis and P. polaris. The evolution of epipelagic calanoids of the Southern Ocean differs in some respects from deepwater pelagic cal- anoids. Throughout the world?s oceans, there are many epi- pelagic marine calanoids whose distribution is confi ned to a water mass or current system within an ocean basin and sometimes very narrowly within that basin. Often, such distributions appear to be restricted to a zone of latitudes, although the causes may be related to specifi c nutrient and temperature regimes (Reid et al., 1978). Epipelagic species from the low or middle latitudes provide many examples of zonally distributed species (Frost and Fleminger, 1968). The circumpolar distributions of the epipelagic endemics of the Southern Ocean, Calanoides acutus, Rhincalanus gigas, Calanus propinquus, Metridia gerlachei, Clauso- calanus laticeps, Calanus simillimus, Clausocalanus bre- vipes, and Ctenocalanus citer, can be interpreted as zonal distributions. The fi rst fi ve are restricted to waters south of the Antarctic Convergence; the last three are restricted to waters south of the Subtropical Convergence and north of the Antarctic Convergence (Table 2). These two sets of epipelagic species may be adapted to the two different zones of cold water bounded by Antarctic Convergence and Subtropical Convergence, as well as by the unique pri- mary productivity of the Southern Ocean. In the Southern Ocean, the evolution of epipelagic calanoids shares some common attributes with deepwa- ter pelagic calanoids. High primary productivity may have enabled the evolution of epipelagic species; all fi ve endemic epipelagic species found south of the Antarctic Convergence are endemic herbivores utilizing the region?s high primary productivity; all are very common. Three of the eight subantarctic epipelagic species are also endemic herbivores and are common or very common. Sibling spe- cies pairs can be found among epipelagic species as well as deepwater species, e.g., Calanoides acutus and C. cari- natus or Rhincalanus gigas and R. nasutus. In contrast to deepwater sibling species pairs, both members of an epi- pelagic pair often are common or very common (Figure 1). Candacia falcifera Calanus australis Chiridiella megadactyla Drepanopus bispinosus Heterorhabdus pustulifer Metridia gerlachei Neocalanus tonsus Paraeuchaeta eltaninae Paraeuchaeta regalis Paraheterorhabdus farrani Scaphocalanus parantarcticus Scaphocalanus farrani Stephos antarcticus C. parafalcifera C. helgolandicus C. abyssalis D. bungei H. norvegica M. longa N. plumchurus P. rubra P. polaris P. longispinus S. acrocephalus S. brevicornis S. arcticus S 35 0 35 N FIGURE 2. Distribution of selected pelagic calanoids of the Southern Ocean and their closest relatives in the subarctic region or the Arctic Ocean. 12_Park_pg143-180_Poles.indd 16712_Park_pg143-180_Poles.indd 167 11/17/08 8:30:24 AM11/17/08 8:30:24 AM 168 SMITHSONIAN AT THE POLES / PARK AND FERRARI Epipelagic, bipolar species pairs have also been identifi ed, e.g., Calanus australis and C. helgolandicus and Neocala- nus tonsus and N. plumchrus. The evolution of coastal species in the Southern Ocean may refl ect processes similar to the model for the deepwater bipolar species pairs. Several species of coastal genera, such as Drepanopus and Stephos, are present in the Southern Ocean and the Arctic Ocean. However, the morphological differences between the Southern Ocean and the Arctic Ocean members are more extensive than those found between members of the oceanic genera of bipolar species pairs. The Southern Ocean coastal species Drepanopus pectinatus, D. bispinosus, and D. forcipatus and Stephos longipes and S. antarcticus are readily distin- guished from their congeners D. bungei and D. furcatus and S. arcticus and its six relatives in the Arctic Ocean. In contrast, the Southern Ocean species of oceanic gen- era are diffi cult to distinguish from their Arctic and boreal congeners because they are very similar morphologically. Apparently, the species of these coastal pelagic genera may have had a different evolutionary history and perhaps a different biogeographical history than the oceanic pelagic calanoid species, although similar processes may have af- fected both groups. High productivity may have played an important role in the structure of the nearshore fauna be- cause fi ve of eight coastal species endemic to the Southern Ocean (Table 1) are common or very common. The situation for Paralabidocera antarctica, P. gran- dispina, and P. separabilis requires further consideration. The genus Paralabidocera, an acartiid restricted to the Southern Hemisphere, is one of only fi ve pelagic marine genera that are limited to one of the two hemispheres. The others are Epilabidocera, a pontellid, Eurytemora, a temorid, Jashnovia, an aetideid, and Pseudocalanus, a clausocalanid, and these four are limited to the Northern Hemisphere. Species of these genera are found in estua- rine or inshore waters or in the neritic zone of the oceans. Each of these genera has a morphologically similar ge- nus, and presumed closest relative (T. Park, unpublished data), distributed broadly throughout the world: species of Paralabidocera are similar to those of Acartia, species of Epilabidocera to Labidocera, Eurytemora to Temora, Jashnovia to Gaetanus, and Pseudocalanus to Clauso- calanus. As a result, it seems reasonable to assume that the species of Paralabidocera have evolved from a cosmo- politan acartiid ancestor. In general, morphological dif- ferences between each limited genus and its cosmopolitan relative are not as great as between each limited genus and its remaining confamilial relatives, so that morphological differences between species of Paralabidocera and Acar- tia are not as great as between Paralabidocera and species of Acartiella and Paracartia. The evolution of Paralabi- docera, then, may be a relatively recent event within the Southern Ocean. In summary, most genera of pelagic marine calanoid copepods found in the Southern Ocean are also found north of the Southern Ocean; Paralabidocera, restricted to the Southern Hemisphere, may be an exception. Many species of pelagic calanoid copepods endemic to the Southern Ocean are common or very common, refl ecting their adaptation to the eutrophic environment there. The closest relative of many Southern Ocean species is usually a widely distributed congener adapted to waters of low primary and secondary productivity whose range does not extend into the Southern Ocean. These observations suggest that Southern Ocean endemics evolved from a common oligotrophic ancestor that split into two popula- tions. The widely distributed daughter species retained its adaptation to oligotrophic habitats; the Southern Ocean endemic daughter became adapted to the eutrophic envi- ronment of the Southern Ocean. The antarctica species group of Paraeuchaeta appears to be monophyletic and may have subsequently evolved from a common South- ern Ocean eutrophic daughter population after its initial split from an ancestor, similar to P. bisinuata, which is associated with oligotrophic habitats. Therefore, most Southern Ocean endemic species appear to have evolved after the Southern Ocean became an area of high pri- mary and secondary productivity. High productivity is assumed to have developed with a strong circumpolar current following the separation of Antarctica from Aus- tralia. A hypothesis structuring the benthopelagic cala- noid fauna from the divergence in feeding appendages differentially adapted for detritivory (Markhaseva and Ferrari, 2006) shares the same phenomenon, adaptation to diversity in food availability as an evolutionary cause of species diversity. Another hypothesis to explain the endemicity of the Southern Ocean involves species that contribute to bipolar species pairs. These may have evolved from a widely dis- tributed deepwater species with shallow polar populations whose intervening deepwater populations subsequently be- came extinct, leaving a species with a bipolar distribution. The two populations then diverged. In this model, bipolar species are transient natural phenomena. Finally, this re- view provides no unequivocal support for zonal distribu- tions for mesopelagic or bathypelagic calanoid species. In general, mesopelagic or bathypelagic calanoids appear to 12_Park_pg143-180_Poles.indd 16812_Park_pg143-180_Poles.indd 168 11/17/08 8:30:26 AM11/17/08 8:30:26 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 169 occur broadly throughout the world?s oceans and are unre- stricted latitudinally or longitudinally. FUTURE CONSIDERATIONS Seventy-seven pelagic, marine calanoid copepods had been reported from the Southern Ocean prior to 1950. Vervoort (1951, 1957) added 26 species, bringing the to- tal number to 103. A total of 117 calanoid species were known before Park (1978, 1980, 1982, 1983a, 1983b, 1988, 1993, 2000) added 73 species. During this period, a few other authors added several species, so that the total reached 201 species by the end of the century. Recently, three species of Metridia and one species of Lucicutia have been added (Markhaseva, 2001; Markhaseva and Ferrari, 2005), bringing the total to 205 species. The midwater trawls employed by the U.S. Antarctic Research Program have been very effective in sampling the large pelagic copepods of the Southern Ocean. There can be very few pelagic species left to be collected with sampling gear of this kind of device. However, fi ne-mesh samples, less than 100 micrometers, taken with traditional sampling gear like the conical plankton net may add new pelagic species or new records of pelagic species already known from other areas of the world?s oceans. With new sampling methods capable of reaching un- explored habitats, more species of calanoids, many ex- pected to be new to science, can be anticipated to increase the Southern Ocean fauna of calanoid copepods. Waters immediately above the seabed, where new benthopelagic calanoid copepods have only recently been collected and studied carefully, are an example. The diverse fauna of this habitat has been very poorly sampled, and lists of new spe- cies and new records are in a growth phase. However, new species and new records of benthopelagic copepods are not expected to effect the conclusions drawn here about pelagic calanoids. Beginning with the studies by Vervoort (1950, 1951, 1957), the species descriptions available in the literature for the Southern Ocean calanoids have been based on the complete morphology of the exoskeleton. Generally, these descriptions are of excellent quality, and the observations are easily repeatable with newly collected specimens. Most of the species discovered earlier have been redescribed in detail by subsequent authors, making the identifi cation of specimens of those species reliable. However, the zoo- geographic distribution of most species, particularly in areas outside of the Southern Ocean, needs signifi cant attention. Information about the vertical distribution of many species, except for the very common ones, remains insuffi cient. Finally, the population structure, particularly for nauplii, remains poorly known. These problems can be addressed with contemporary sampling gear but will require the kind of intellectual curiosity which drove the early exploration of the Southern Ocean by the U.S. Ant- arctic Research Program. ACKNOWLEDGMENTS We would like to thank former Under Secretary for Science Dr. David Evans and the Offi ce of the Under Sec- retary for Science at the Smithsonian Institution for spon- soring the International Polar Year Symposium on 3 and 4 May 2007 in Washington, D.C. Rafael Lemaitre, Chair- man of the Department of Invertebrate Zoology, kindly invited one of us (TP) to participate in the symposium. 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APPENDIX 1 This appendix contains a list of pelagic calanoid cope- pod species, alphabetical by genus and then species, reported from the Southern Ocean. Each species name includes the author and date of publication of the original species de- scription. The author and date entry listed below a species name is the latest taxonomic reference to the species. Ab- breviations for distributions are as follows: Ant, Antarctic; sAnt, subantarctic; Stemp, south temperate; Trop, tropical; Ntemp, north temperate; sArc, subarctic; Arc, Arctic. Aetideopsis antarctica (Wolfenden, 1908): Ant Faroella antarctica Wolfenden, 1908:39, pl. 2, fi gs. 1? 4. Aetideopsis antarctica Bradford, 1971:18, fi gs. 31? 48. Aetideopsis minor (Wolfenden, 1911): Ant, Arc Faroella minor Wolfenden, 1911:214. Aetideopsis minor Park, 1978:115? 118, fi gs. 8? 9; Markhaseva, 1996:32, fi gs. 20? 21. Aetideopsis multiserrata (Wolfenden, 1904): Ant, sArc Faroella multiserrata Wolfenden, 1904:117, pl. 9, fi gs. 26? 28. Aetideopsis multiserrata Park, 1978:111? 115, fi gs. 6? 7; Markhaseva, 1996:37, fi gs. 22? 23. Aetideopsis rostrata Sars, 1903: Ant, sArc, Arc Aetideopsis rostrata Sars, 1903:160, pls. 4, 5; Markhaseva, 1996:42, fi gs. 26? 28. Aetideopsis infl ata Park, 1978:118? 122, fi gs. 11? 12. Aetideopsis tumorosa Bradford, 1969: sAnt Aetideopsis tumorosa Bradford, 1969:74, fi gs. 1? 28; Park, 1978:118, fi g. 10; Markhaseva, 1996:42, fi g. 29. Aetideus arcuatus (Vervoort, 1949): sAnt, Ntemp Snelliaetideus arcuatus Vervoort, 1949:4, fi g. 1; Park, 1978:108? 111, fi gs. 4? 5. Aetideus arcuatus Markhaseva, 1996:14, fi g. 3. Aetideus australis (Vervoort, 1957): Ant, sAnt Euaetideus australis Vervoort, 1957:46, fi gs. 16? 19, 20a; Park, 1978:105? 108, fi gs. 2? 3. Arietellus simplex Sars, 1905: Ant, sArc Arietellus simplex Sars, 1905:5, 22; 1925:334, pl. 120, fi gs. 7? 12; Vervoort, 1957:141. Augaptilus glacialis Sars, 1900: Ant, Arc Augaptilus glacialis Sars, 1900:88, pl.s 26, 27; 1925:254, pl. 76, fi gs.1? 6; Vervoort, 1951:144, fi gs. 80, 81. Batheuchaeta antarctica Markhaseva, 1986: Ant Batheuchaeta antarctica Markhaseva, 1986:848, fi g. 6; 1996:53, fi g. 34. 12_Park_pg143-180_Poles.indd 17312_Park_pg143-180_Poles.indd 173 11/17/08 8:30:30 AM11/17/08 8:30:30 AM 174 SMITHSONIAN AT THE POLES / PARK AND FERRARI Batheuchaeta lamellata Brodsky, 1950: Ant-Ntemp Batheuchaeta lamellata Brodsky, 1950:189, fi gs. 106? 107; Markhaseva, 1996:57? 58. Batheuchaeta peculiaris Markhaseva, 1983: Ant-Ntemp Batheuchaeta peculiaris Markhaseva, 1983:1740, fi g. 1; 1996:58, fi gs. 41? 42. Batheuchaeta pubescens Markhaseva, 1986: Ant Batheuchaeta pubescens Markhaseva, 1986:846, fi g. 5; 1996:58, fi g. 43. Bathycalanus bradyi (Wolfenden, 1905): Ant-Ntemp Megacalanus bradyi Wolfenden, 1905:1? 3, pl. 1, fi gs. 1? 6. Bathycalanus maximus Wolfenden, 1911:198, pl. 23, fi gs. 1? 7, text fi g. 2. Bathycalanus bradyi Vervoort, 1957:32, fi g. 7. Bathycalanus eltaninae Bj?rnberg, 1968: sAnt Bathycalanus eltaninae Bj?rnberg, 1968:75, fi gs. 15? 41. Bathycalanus infl atus Bj?rnberg, 1968: sAnt Bathycalanus infl atus Bj?rnberg, 1968:81, fi gs. 42? 54. Bathycalanus unicornis Bj?rnberg, 1968: sAnt Bathycalanus unicornis Bj?rnberg, 1968:73, fi gs. 1? 14. Bradycalanus enormis Bj?rnberg, 1968: sAnt Bradycalanus enormis Bj?rnberg, 1968:85, fi gs. 64? 77. Bradycalanus pseudotypicus Bj?rnberg, 1968: sAnt Bradycalanus pseudotypicus Bj?rnberg, 1968:82, fi gs. 55? 63, 78. Calanoides acutus (Giesbrecht, 1902): Ant Calanoides acutus Giesbrecht, 1902:17, pl. 1, fi gs. 10? 14; Vervoort, 1951:42, fi gs. 25? 33. Calanus australis Brodsky, 1959: sAnt-Stemp Calanus australis Brodsky, 1959:1539? 1542, pl. 1, fi gs. 9? 12; pl. 2, fi gs. 4, 8, 10, 11; pl. 3, fi gs. 11, 13, 14; pl. 4, fi gs. 8, 9. Calanus propinquus Brady, 1883: Ant Calanus propinquus Brady, 1883:34, pl. 2, fi gs. 1? 7; pl. 14, fi gs. 10? 11; Vervoort, 1951:27, fi gs. 14? 24. Calanus simillimus Giesbrecht, 1902: sAnt Calanus simillimus Giesbrecht, 1902:16, fi g. 9; Vervoort, 1951:11, fi gs. 3? 14. Candacia cheirura Cleve, 1904: sAnt Candacia cheirura Cleve, 1904:180, 186, 198, pl. 1, fi gs. 1? 6; pl. 2, fi gs. 7? 10; Farran, 1929:273, fi g. 29; Vervoort, 1957:142. Candacia falcifera Farran, 1929: Ant-sArc Candacia falcifera Farran, 1929:270, fi g. 28; Vervoort, 1957:142, fi g. 132. Candacia maxima Vervoort, 1957: Ant-sAnt Candacia maxima Vervoort, 1957:142? 144, fi gs.132? 138. Centraugaptilus rattrayi (Scott, 1894): sAnt-sArc Augapatilus rattrayi Scott, 1894:36, pl. 2, fi gs. 25? 37. Centraugaptilus rattrayi Sars, 1925:304, pl. 106; Hardy and Gun- ther, 1935:183. Cephalophanes frigidus Wolfenden, 1911: Ant-sAnt Cephalophanes frigidus Wolfenden, 1911:284? 285, fi g. 46; Park, 1983b:321? 325, fi gs. 3? 4. Chiridiella megadactyla Bradford, 1971: Ant Chiridiella megadactyla Bradford, 1971:19? 20, fi gs. 49? 60. Chiridiella subaequalis Grice and Hulsemann, 1965: Ant-Ntemp Chiridiella subaequalis Grice and Hulsemann, 1965:231? 235, fi g. 10a? l; Markhaseva, 1996:108, fi g. 82. Chiridius gracilis Farran, 1908: Ant-sArc Chiridius gracilis Farran, 1908:30, pl. 2, fi gs. 1? 3; Park, 1978:122? 124, fi g. 13; Markhaseva, 1996:111, fi gs. 83? 84. Chiridius polaris Wolfenden, 1911: Ant, Ntemp Chiridius polaris Wolfenden, 1911:211, text fi g. 6, pl. 24, fi gs. 9? 12; Markhaseva, 1996:119, fi gs. 94? 97. Chiridius subantarcticus Park, 1978:125? 127, fi g. 14. Chirundina streetsii Giesbrecht, 1895: sAnt-sArc Chirundina streetsii Giesbrecht, 1895:249, pl. 1, fi gs. 5? 10; Park, 1978:179, fi gs. 52? 53. Clausocalanus brevipes Frost and Fleminger, 1968: sAnt Clausocalanus brevipes Frost and Fleminger, 1968:70, fi gs. 56? 59. Clausocalanus brevipes Farran, 1929: Ant Clausocalanus laticeps Farran, 1929:224, fi g. 4; Frost and Flem- inger, 1968:42, fi gs. 24? 28. Cornucalanus chelifer (Thompson, 1903): Ant-Ntemp Scolecithrix chelifer Thompson, 1903:21, pl. 5, fi gs. 1? 9; Park, 1983b:352? 357, fi gs. 23? 26. Cornucalanus robustus Vervoort, 1957: Ant-Trop Cornucalanus robustus Vervoort, 1957:88? 91, fi gs. 71? 76; Park, 1983:358? 363, fi gs. 27? 30. Cornucalanus simplex Wolfenden, 1905: Ant-Ntemp Cornucalanus simplex Wolfenden, 1905:22; Park, 1983b:364? 365, fi g. 31. Ctenocalanus citer Heron and Bowman, 1971: sAnt Ctenocalanus citer Heron and Bowman, 1971:142, fi gs. 1, 16? 18, 31? 36, 54? 58, 71? 77, 94? 99, 130? 150. Ctenocalanus vanus Vervoort, 1951:59? 61; 1957:37. Disseta palumbii Giesbrecht, 1889: sAnt-sArc Disseta palumbii Giesbrecht, 1889a:812; 1893:369, pl. 29, fi gs. 2, 8, 14, 19, 23? 25, 27; pl. 38, fi g. 44; Park, 2000:14? 18, fi gs. 1? 3. Drepanopus bispinosus Bayly, 1982; Ant Drepanopus bispinosus Bayly, 1982:165, fi gs. 2a? 2h, 3a? 3f. Drepanopus forcipatus Giesbrecht, 1888: sAnt Drepanopus forcipatus Giesbrecht, 1888:335; Hulsemann, 1985: 911, fi gs. 2? 4, 6? 8, 10, 12, 14, 16? 20, 23, 24, 27? 29, 32? 33. Drepanopus pectinatus Brady, 1883: sAnt Drepanopus pectinatus Brady, 1883:77, pl. 24, fi gs. 1? 11; Hulsemann, 1985a:910, fi gs. 1, 5, 9, 11, 13, 15, 21, 22, 25, 26, 30, 31. Euaugaptilus aliquantus Park, 1993: sAnt-Stemp Euaugaptilus aliquantus Park, 1993:13? 14, fi gs. 7? 8. Euaugaptilus angustus (Sars, 1905): sAnt-Ntemp Augaptilus angustus Sars, 1905:10? 11. Euaugaptilus angustus Sars, 1925:281? 282, pl. 91; Park, 1993:27? 30, fi gs. 19? 20. Euaugaptilus antarcticus Wolfenden, 1911: Ant Euaugaptilus antarcticus Wolfenden, 1911:334? 336, fi g. 70, pl. 36, fi gs. 6? 7; Park, 1993:32? 37, fi gs. 23? 25. Euaugaptilus austrinus Park, 1993: Ant Euaugaptilus austrinus Park, 1993:37? 41, fi gs. 26? 28. Euaugaptilus brevirostratus Park, 1993: sAnt-Stemp Euaugaptilus brevirostratus Park, 1993:19? 22, fi gs. 12? 14. Euaugaptilus bullifer (Giesbrecht, 1889): Ant-Ntemp Augaptilus bullifer Giesbrecht, 1889a:813; 1893:400, pl. 28, fi gs. 6, 21, 24; pl. 39, fi g. 46. Euaugaptilus bullifer (Giesbrecht, 1889); Park, 1993:22? 25, fi gs. 15? 16. Euaugaptilus gibbus (Wolfenden, 1904): sAnt-Ntemp Augaptilus gibbus Wolfenden, 1904:122; 1911:337? 339, fi g. 72, pl. 37, fi gs. 2? 3. Euaugaptilus gibbus (Wolfenden, 1904); Park, 1993:25? 27, fi gs. 17? 18. Euaugaptilus hadrocephalus Park, 1993: Ant-Stemp Euaugaptilus hadrocephalus Park, 1993:6? 9, fi gs. 3, 4. Euaugaptilus laticeps (Sars, 1905): sAnt-Ntemp Augaptilus laticeps Sars, 1905b:11. Euaugaptilus laticeps Sars, 1925:264? 265, pl. 80; Park, 1993:30? 32, fi gs. 21? 22. Euaugaptilus magnus (Wolfenden, 1904): Ant-Ntemp Augaptilus magnus Wolfenden, 1904:111, 122, 142, 145; Wolfenden 1911:341? 343, fi g. 74, pl. 37, fi gs. 4? 9. Euaugaptilus magnus Park, 1993:41? 44, fi gs. 29? 30. Euaugaptilus maxillaris Sars, 1920: Ant-Ntemp Euaugaptilus maxillaris Sars, 1920:15; 1925:287? 288, pl. 95; Park, 1993:5? 6, fi gs. 1? 2. 12_Park_pg143-180_Poles.indd 17412_Park_pg143-180_Poles.indd 174 11/17/08 8:30:30 AM11/17/08 8:30:30 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 175 Euaugaptilus nodifrons (Sars, 1905): Ant-Ntemp Augaptilus nodifrons Sars, 1905b:13? 14. Euaugaptilus nodifrons Sars, 1925:267? 269, pl. 82; Park, 1993:14? 19, fi gs. 9? 11. Euaugaptilus oblongus (Sars, 1905): sAnt-Ntemp Augaptilus oblongus Sars, 1905b:11. Euaugaptilus oblongus Sars, 1925:266? 267, pl. 81; Park, 1993:44? 47, fi gs. 31? 32. Euaugaptilus perasetosus Park, 1993: Ant-Stemp Euaugaptilus perasetosus Park, 1993:9? 13, fi gs. 5? 6. Eucalanus hyalinus (Claus, 1866): sAnt-Ntemp Calanella hyaline Claus, 1866:8. Eucalanus hyalinus Bradford? Grieve, 1994:76, fi gs. 42, 88. Euchirella maxima Wolfenden, 1905: sAnt-sArc Euchirella maxima Wolfenden, 1905:18, pl. 6, fi gs. 9? 11; Park, 1978:149? 151, fi g. 30. Euchirella rostrata (Claus, 1866): sAnt-Ntemp Undina rostrata Claus, 1866:11, pl. 1, fi g. 2. Euchirella rostrata Park, 1978:147? 149, fi g. 29. Euchirella rostromagna Wolfenden, 1911: Ant Euchirella rostromagna Wolfenden, 1911:235; Park, 1978:151? 155, fi gs. 31? 34. Euchirella similis Wolfenden, 1911: sAnt-Trop Euchirella similis Wolfenden, 1911:238, text fi g. 23, pl. 28, fi gs. 1? 2; Park, 1978:155? 158, fi gs. 35? 36. Farrania frigida (Wolfenden, 1911): Ant-Trop Drepanopsis frigida Wolfenden, 1911:245, text fi g. 29;Vervoort, 1951:61, fi gs. 34? 39. Farrania frigida Vervoort, 1957:38? 39. Gaetanus antarcticus Wolfenden, 1905: Ant-Ntemp Gaetanus antarcticus Wolfenden, 1905:7, pl. 3, fi g. 1; Park, 1978: 141? 144, fi gs. 25, 26; Markhaseva, 1996:178, fi gs. 138? 139. Gaetanus brevispinus (Sars, 1900): Ant-Arc Chiridius brevispinus Sars, 1900:68, pl. 19; Markhaseva, 1996:187? 195, fi gs. 149? 152. Gaidius intermedius Wolfenden, 1905:6, pl. 3, fi gs. 4? 5; Park, 1978:131? 136, fi gs. 18? 20. Gaetanus kruppii Giesbrecht, 1903: sAnt-sArc Gaetanus kruppii Giesbrecht, 1903:22; Park, 1978:136? 139, fi gs. 21? 22; Markhaseva, 1996:196? 201, fi gs. 157? 158. Gaetanus latifrons Sars, 1905: sAnt-sArc Gaetanus latifrons Sars, 1905a: 4, 11; Vervoort, 1957:61? 62; Markhaseva, 1996, p.201? 204, fi gs. 159? 160. Gaetanus minor Farran, 1905: sAnt-Ntemp Gaetanus minor Farran, 1905:34, pl. 5; Park, 1978:144? 147, fi gs. 27? 28; Markhaseva, 1996:205? 206, fi g. 164. Gaetanus paracurvicornis Brodsky, 1950: Ant-Ntemp Gaetanus paracurvicornis Brodsky, 1950:167, fi g. 84; Markhaseva, 1996:211, fi g. 167. Gaetanus pileatus Farran, 1903: sAnt-Ntemp Gaetanus pileatus Farran, 1903:16, pl. 17, fi gs. 1? 11; Park, 1978:139? 141, fi gs. 23? 24; Markhaseva, 1996:211? 212, fi gs. 168? 169. Gaetanus tenuispinus (Sars, 1900): Ant-Arc Chiridius tenuispinus Sars, 1900:67, pl. 18. Gaidius tenuispinus Park, 1978, pp.127? 131, fi gs. 15? 17. Gaetanus tenuispinus Markhaseva, 1996:221? 225, fi gs. 177? 178. Haloptilus fons Farran, 1908: Ant-Ntemp Haloptilus fons Farran, 1908:69? 71, pl. 7, fi gs. 11? 15; Park, 1988:3? 4, fi gs. 1? 2. Haloptilus longicirrus Brodsky, 1950: Ant-sArc Haloptilus longicirrus Brodsky, 1950:363? 364, fi g. 254; Park, 1988:21? 22, fi g. 13. Haloptilus ocellatus Wolfenden, 1905: Ant Haloptilus ocellatus Wolfenden, 1905:14? 15, pl. 5; Park, 1988:4? 9, fi gs. 3? 4. Haloptilus oxycephalus (Giesbrecht, 1889): Ant-Ntemp Hemicalanus oxycephalus Giesbrecht, 1889a:813; Giesbrecht, 1893: 384, pl. 42, fi gs. 16, 23. Haloptilus oxycephalus Park, 1988:9? 14, fi gs. 5? 6. Heterorhabdus austrinus Giesbrecht, 1902: Ant-sAnt Heterorhabdus austrinus Giesbrecht, 1902:28, pl. 6, fi gs. 1? 9; Park, 2000:132? 133, fi gs. 104? 105. Heterorhabdus lobatus Bradford, 1971: sAnt-Ntemp Heterorhabdus lobatus Bradford, 1971:131, fi gs. 9, 10a? c; Park, 2000:105? 106, fi g. 74. Heterorhabdus paraspinosus Park, 2000: sAnt-Stemp Heterorhabdus paraspinosus Park, 2000:131? 132, fi gs. 102? 103. Heterorhabdus pustulifer Farran, 1929: Ant-sAnt Heterorhabdus pustulifer Farran, 1929:266, fi g. 27; Park, 2000: 124? 125, fi gs. 92? 93. Heterorhabdus spinosus Bradford, 1971: sAnt-Stemp Heterorhabdus spinosus Bradford, 1971:121, fi gs.1, 2g? k, 3c, 4c; Park, 2000:130? 131, fi gs. 100? 101. Heterostylites nigrotinctus Brady, 1918: Ant-sAnt Heterostylites nigrotinctus Brady, 1918:27, pl. 6, fi gs. 1? 8; Park, 2000:44? 45, fi gs. 22? 23. Landrumius antarcticus Park, 1983a: Ant Landrumius antarcticus Park, 1983a:192? 195, fi gs. 15? 16. Landrumius gigas (Scott, 1909): sAnt-Trop Brachycalanus gigas Scott, 1909:81? 82, pl. 35, fi gs. 10? 18. Lophothrix gigas Grice and Hulsemann, 1968:332? 334, fi gs. 63? 74. Landrumius gigas Park, 1983:195? 197, fi gs. 17? 18. Lophothrix frontalis Giesbrecht, 1895: sAnt-Ntemp Lophothrix frontalis Giesbrecht, 1895:254? 255, pl. 2, fi gs. 1? 5, 9? 12; Park, 1983a:178? 184, fi gs. 7? 10. Lophothrix humilifrons Sars, 1905: Ant-Ntemp Lophothrix humilifrons Sars, 1905a:22; 1925:166? 167, pl. 46, fi gs. 15? 22; Park, 1983a:184? 188, fi gs. 11? 12. Lucicutia bradyana Cleve, 1904: Ant-Trop Lucicutia bradyana Cleve, 1904:204? 206, pl. 6, fi gs. 33, 34; Markhaseva and Ferrari, 2005:1084? 1091, fi gs. 3? 7. Lucicutia curta Farran, 1905: Ant-sArc Lucicutia curta Farran, 1905:44, pl. 12, fi gs. 1? 7; Vervoort, 1957:128? 129, fi gs. 114? 117. Lucicutia macrocera Sars, 1920: Ant-sArc Lucicutia macrocera Sars, 1920:10; 1925:213, pl. 57, fi gs. 12? 15; Vervoort, 1957:130, fi gs. 118, 119. Lucicutia magna Wolfenden, 1903: Ant-sArc Lucicutia magna Wolfenden, 1903:124; Wolfenden, 1911:316? 317, text fi g. 59; Hulsemann, 1966:727, fi g. 119. Lucicutia ovalis (Giesbrecht, 1889): Ant-sArc Isochaeta ovalis Giesbrecht, 1889a:812. Lucicutia frigida Wolfenden, 1911:320, text fi g. 62; Vervoort, 1957:126? 128, fi gs. 111? 114. Lucicutia wolfendeni Sewell, 1932: Ant-sArc Lucicutia wolfendeni Sewell, 1932:289; Markhaseva and Ferrari, 2005:1091? 1094, fi gs. 8? 9. Megacalanus princeps Wolfenden, 1904: Ant-sArc Megacalanus princeps Wolfenden, 1904:49, fi g. 4; Vervoort, 1957: 32, fi g. 7. Metridia brevicauda Giesbrecht, 1889: sAnt-sArc Metridia brevicauda Giesbrecht, 1889b:24; 1893:340, pl. 33, fi gs. 5, 10? 11, 14, 21, 32; Vervoort, 1957:122. Metridia curticauda Giesbrecht, 1889: Ant-sArc Metridia curticauda Giesbrecht, 1889b:24; 1893:340, pl. 33, fi gs. 4, 15, 33; Vervoort, 1951:121, fi gs. 65? 67; 1957:122. Metridia ferrarii Markhaseva, 2001: Ant-Ntemp Metridia ferrarii Markhaseva, 2001:44? 46, fi gs. 1? 59. Metridia gerlachei Giesbrecht, 1902: Ant Metridia gerlachei Giesbrecht, 1902:27, pl. 5, fi gs. 6? 14; Vervoort, 1951:120; 1957:120? 121, fi gs. 109, 110. 12_Park_pg143-180_Poles.indd 17512_Park_pg143-180_Poles.indd 175 11/17/08 8:30:31 AM11/17/08 8:30:31 AM 176 SMITHSONIAN AT THE POLES / PARK AND FERRARI Metridia lucens Boeck, 1864: sAnt-Ntemp Metridia lucens Boeck, 1864:238; Vervoort, 1957:119. Metridia hibernica Giesbrecht, 1893:345, pl. 32, fi g. 11; pl. 35, fi gs. 2, 12, 16, 22, 28, 36, 39. Metridia ornata Brodsky, 1950: Ant-sArc Metridia ornata Brodsky, 1950:303? 305, fi g. 210; Markhaseva, 2001:48? 49, fi gs. 184? 243. Metridia princeps Giesbrecht, 1889: Ant-sArc Metridia princeps Giesbrecht, 1889b:24; Markhaseva, 2001:47? 48, fi gs. 148? 183. Metridia pseudoasymmetrica Markhaseva, 2001: Ant-sAnt Metridia pseudoasymmetrica Markhaseva, 2001:46? 47, fi gs. 60? 110. Metridia venusta Giesbrecht, 1889: sAnt-Ntemp Metridia venusta Giesbrecht, 1889b:24; 1893:340, pl. 33, fi gs. 7, 17, 29; Vervoort, 1957:121? 122. Microcalanus pygmaeus (Sars, 1900): Ant-Arc Pseudocalanus pygmaeus Sars, 1900:73, pl. 21. Microcalanus pygmaeus Vervoort, 1957:36? 37, fi g. 9; Bradford- Grieve, 1994:130, fi g. 75. Mimocalanus cultrifer Farran, 1908: Ant-sArc Mimocalanus cultrifer Farran, 1908:23, pl. 1, fi gs. 5? 9; Damkaer, 1975:68? 69, fi gs. 164? 168. Neocalanus tonsus (Brady, 1883): sAnt-Stemp Calanus tonsus Brady, 1883:34, pl. 4, fi gs. 8, 9; Vervoort, 1957:27, fi gs. 3? 6. Neocalanus tonsus Bradford-Grieve, 1994:42, fi gs. 17, 82. Nullosetigera bidentatus (Brady, 1883): Ant-sArc Phyllopus bidentatus Brady, 1883:78, pl. 5, fi gs. 7? 16; Vervoort, 1957:141. Onchocalanus cristatus (Wolfenden, 1904): Ant-Ntemp Xanthocalanus cristatus Wolfenden, 1904:119, pl. 9, fi gs. 18? 19. Onchocalanus cristatus Park, 1983b:335? 343, fi gs. 13? 15. Onchocalanus hirtipes Sars, 1905: Ant-Ntemp Onchocalanus hirtipes Sars, 1905a:20? 21; 1925:148? 149, pl. 41, fi gs. 6? 11; Park, 1983b:351, fi g. 22. Onchocalanus magnus (Wolfenden, 1906): Ant Xanthoclanus magnus Wolfenden, 1906:32? 33, pl. 10, fi gs. 7? 9. Onchocalanus magnus Park, 1983:343? 347, fi gs. 16? 19. Onchocalanus paratrigoniceps Park, 1983b: Ant-Stemp Onchocalanus paratrigoniceps Park, 1983b:333? 335, fi gs. 10? 12. Onchocalanus subcristatus (Wolfenden, 1906): Ant Xanthocalanus subcristatus Wolfenden, 1906:31? 32, pl. 10, fi gs. 4? 6. Onchocalanus subcristatus Wolfenden, 1911:278, pl. 31, fi gs. 6? 8. Onchocalanus trigoniceps Sars, 1905: Ant-Ntemp Onchocalanus trigoniceps Sars, 1905a:20; 1925:144? 147, pl. 40, fi gs. 1? 15; Park, 1983b:329? 333, fi gs. 7? 9. Onchocalanus wolfendeni Vervoort, 1950: Ant Onchocalanus wolfendeni Vervoort, 1950:22? 26, fi gs. 9? 11; Park, 1983b:347? 351, fi gs. 20? 21. Pachyptilus eurygnathus Sars, 1920: Ant-sArc Pachyptilus eurygnathus Sars, 1920:18; 1925:321, pl. 114; Vervoort, 1957:140. Paraeuchaeta abbreviata (Park, 1978): Ant-Trop Euchaeta abbreviata Park, 1978:240? 244, fi gs. 92, 93. Paraeuchaeta abbreviata Park, 1995:63? 64, fi gs. 58? 59. Paraeuchaeta antarctica (Giesbrecht, 1902): Ant-sAnt Euchaeta antarctica Giesbrecht, 1902:21, pl. 3, fi gs. 1? 8; Fontaine, 1988:32? 38, fi gs. 3? 8. Paraeuchaeta antarctica Park, 1995:88? 89, fi gs. 84? 85. Paraeuchaeta austrina (Giesbrecht, 1902): Ant Euchaeta austrina Giesbrecht, 1902:22, pl. 3, fi gs. 9? 16; Fontaine, 1988:46? 49, fi gs. 6, 13, 16, 17. Paraeuchaeta barbata (Brady, 1883): Ant-Arc Euchaeta barbata Brady, 1883:66, pl. 22, fi gs. 6? 12. Paraeuchaeta barbata Park, 1995:37? 38, fi g. 23. Paraeuchaeta biloba Farran, 1929: Ant-sAnt Paraeuchaeta biloba Farran, 1929:242, fi g. 11. Euchaeta biloba Park, 1978:217? 220, fi gs. 74? 76. Paraeuchaeta comosa Tanaka, 1958: sAnt-Ntemp Paraeuchaeta comosa Tanaka, 1958:363, fi g. 79a? g; Park, 1995:56? 57, fi g. 50. Paraeuchaeta dactylifera (Park, 1978) Ant-sAnt Euchaeta dactylifera Park, 1978:240, fi g. 91. Paraeuchaeta dactylifera Park, 1995:70, fi g. 67. Paraeuchaeta eltaninae (Park, 1978): Ant Euchaeta eltaninae Park, 1978:280? 283, fi gs. 118? 120. Paraeuchaeta eltaninae Park, 1995:39, fi g. 25. Paraeuchaeta erebi Farran, 1929: Ant Paraeuchaeta erebi Farran, 1929:239, fi g. 9. Euchaeta erebi Fontaine, 1988:41? 45, fi gs. 11? 13. Paraeuchaeta exigua (Wolfenden, 1911): sAnt-Stemp Euchaeta exigua Wolfenden, 1911:300, text fi g. 52. Paraeuchaeta exigua Park, 1995:77? 78, fi g. 74. Paraeuchaeta hansenii (With, 1915): sAnt-sArc Euchaeta hansenii With, 1915:181, text fi g. 52. Paraeuchaeta hansenii Park, 1995:57? 58, fi g. 51. Paraeuchaeta kurilensis Heptner, 1971: Ant-sArc Paraeuchaeta kurilensis Heptner, 1971:83, fi g. 4; Park, 1995:62? 63, fi g. 57. Paraeuchaeta parvula (Park, 1978): Ant-sAnt Euchaeta parvula Park, 1978:256? 259, fi gs. 102? 104. Paraeuchaeta parvula Park, 1995:38, fi g. 24. Paraeuchaeta pseudotonsa (Fontaine, 1967): sAnt-Ntemp Euchaeta pseudotonsa Fontaine, 1967:204, fi gs. 1B, 2B, 3B, 3E, 6B, 6E, 7B, 8B, E, 9A, 9C, 10, 12. Paraeuchaeta pseudotonsa Park, 1995:74? 75, fi g. 71. Paraeuchaeta rasa Farran, 1929: Ant-sAnt Paraeuchaeta rasa Farran, 1929:240, fi g. 10; Park, 1995:46, fi g. 35. Paraeuchaeta regalis (Grice and Hulsemann, 1968): Ant-Stemp Euchaeta regalis Grice and Hulsemann, 1968:329, fi gs. 34? 40. Paraeuchaeta regalis Park, 1995:50, fi g. 41. Paraeuchaeta sarsi (Farran, 1908): sAnt-Ntemp Euchaeta sarsi Farran, 1908:41, pl. 3, fi gs. 15? 16. Paraeuchaeta sarsi Park, 1995:47? 48, fi gs. 37? 38. Paraeuchaeta scotti (Farran, 1908): sAnt-Ntemp Euchaeta scotti Farran, 1908:42, pl. 3, fi gs. 11? 12. Paraeuchaeta scotti Park, 1995:40? 41, fi g. 27. Paraeuchaeta similis (Wolfenden, 1908): Ant Euchaeta similis Wolfenden, 1908:19, pl. 4, fi gs. 1? 4; Park, 1978:227? 229, fi g. 82; Fontaine, 1988:38? 41, fi gs. 6, 9, 10. Paraeuchaeta tumidula (Sars, 1905): Ant-sArc Euchaeta tumidula Sars, 1905a:15. Paraeuchaeta tumidula Park, 1995:82? 83, fi g. 79. Euchaeta biconvexa Park, 1978:264? 267, fi gs. 108, 109. Paraeuchaeta tycodesma (Park, 1978): Ant Euchaeta tycodesma Park, 1978:229? 231, fi g. 83; Fontaine, 1988: 45? 46, fi gs. 13? 15. Paraheterorhabdus compactus (Sars, 1900): sAnt-Arc Heterorhabdus compactus Sars, 1900:83, pls. 24, 25. Paraheterorhabdus compactus Park, 2000:85? 88, fi gs. 56? 58. Paraheterorhabdus farrani (Brady, 1918): Ant-sAnt Heterorhabdus farrani Brady, 1918:27, pl. 4, fi gs.10? 18. Paraheterorhabdus farrani Park, 2000:78? 80, fi gs. 48, 49. Paralabidocera antarctica (Thompson, 1898): Ant Paracartia antarctica Thompson, 1898:295, pl. 18, fi gs. 1? 12. 12_Park_pg143-180_Poles.indd 17612_Park_pg143-180_Poles.indd 176 11/17/08 8:30:32 AM11/17/08 8:30:32 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 177 Paralabidocera hodgsoni Wolfenden, 1908:26, pl. 6, fi gs. 1? 13. Paralabidocera antarctica Farran, 1929:280; Vervoort, 1951:148. Paralabidocera grandispina Waghorn, 1979: Ant Paralabidocera grandispina Waghorn, 1979:465, fi gs. 5? 8. Paralabidocera separabilis Brodsky and Zvereva, 1976: Ant Paralabidocera separabilis Brodsky and Zvereva, 1976:234, fi gs. 1? 4. Pleuromamma abdominalis (Lubbock, 1856): sAnt-Ntemp Diaptomus abdominalis Lubbock, 1856:22, pl. 10. Pleuromamma abdominalis Steuer, 1932:9? 17, kartes 3? 7; Ver- voort, 1957:123? 124. Pleuromamma antarctica Steuer, 1931: Ant-sAnt Pleuromamma robusta forma antarctica Steuer, 1932:24? 25, karte 10; Vervoort, 1951:123? 126, fi gs. 68, 69; Vervoort, 1957:125. Pleuromamma antarctica Ferrari and Saltzman, 1998:217? 220, fi g. 8. Pleuromamma peseki Farran, 1929: sAnt-Ntemp Pleuromamma gracilis forma peseki Steuer, 1932:34? 36. Pleuromamma peseki Farran, 1929:260, fi gs. 23, 24; Vervoort, 1957:124. Pleuromamma quadrungulata (Dahl, 1893): sAnt-Ntemp Pleuromma quadrungulata Dahl, 1893:105. Pleuromamma quadrungulata Steuer, 1932:26? 30, karte 12? 13; Vervoort, 1957:124. Pleuromamma xiphias (Giesbrecht, 1889): sAnt-Ntemp Pleuromma xiphias Giesbrecht, 1889b:25; 1893:347, pl. 32, fi g. 14; pl. 33, fi gs. 42, 45, 50. Pleuromamma xiphias Steuer, 1932:1? 9, karte 1? 2; Vervoort, 1957:124. Pseudaugaptilus longiremis Sars, 1907: Ant-Arc Pseudaugaptilus longiremis Sars, 1907:24; 1925:310, pl. 109; Ver- voort, 1951:144? 147, fi g. 82; 1957:140. Pseudeuchaeta brevicauda Sars, 1905: Ant-sArc Pseudeuchaeta brevicauda Sars, 1905a:5, 18; Park, 1978:187? 191, fi gs. 57, 58. Pseudochirella batillipa Park, 1978: Ant, Ntemp, Arc Pseudochirella batillipa Park, 1978:176, fi gs. 50, 51; Markhaseva, 1996:255? 256, fi g. 203. Pseudochirella dubia (Sars, 1905): Ant-sArc Undeuchaeta dubia Sars, 1905a:15. Pseudochirella dubia Markhaseva, 1996:262, fi gs. 208? 209. Pseudochirella formosa Markhaseva, 1989: Ant Pseudochirella formosa Markhaseva, 1989:33, fi gs. 1, 7; 1996:264, fi g. 212. Pseudochirella hirsuta (Wolfenden, 1905): Ant-Stemp Euchirella hirsuta Wolfenden, 1905:17, pl. 6, fi gs. 7? 8. Pseudochirella hirsuta Park, 1978:163? 165, fi gs. 41, 42; Markhaseva, 1996:266, fi gs. 214? 215. Pseudochirella mawsoni Vervoort, 1957: Ant-sAnt Pseudochirella mawsoni Vervoort, 1957:64, fi gs. 44? 48; Park, 1978:172? 176, fi gs. 48, 49; Markhaseva, 1996:272? 275, fi gs. 219? 220. Pseudochirella notacantha (Sars, 1905): Ant-sArc Gaidius notacanthus Sars, 1905a:9. Pseudochirella notacantha Markhaseva, 1996:275? 276, fi gs. 221? 222. Pseudochirella obtusa (Sars, 1905): Ant-sArc Undeuchaeta obtusa Sars, 1905a:4, 13. Pseudochirella obtusa Markhaseva, 1996:278, fi gs. 225? 226. Pseudochirella polyspina Park, 1978:169? 172, fi gs. 45? 47. Pseudochirella pustulifera (Sars, 1905): Ant-sArc Undeuchaeta pustulifera Sars, 1905a:14. Pseudochirella pustulifera Park, 1978:165? 169, fi gs. 43? 44. Pseudochirella spectabilis (Sars, 1900): Ant, Arc Undeuchaeta spectabilis Sars, 1900:59, pls. 15, 16. Pseudochirella spectabilis Markhaseva, 1996:289? 290, fi gs. 233? 235. Euchirella elongata Wolfenden, 1905:19, pl. 6, fi gs. 12? 13. Pseudochirella elongata Park, 1978:159? 163, fi gs. 37? 40. Racovitzanus antarcticus Giesbrecht, 1902: Ant, sAnt, sArc Racovitzanus antarcticus Giesbrecht, 1902:26? 27, pl. 4, fi gs. 8? 13; pl. 5, fi gs.1? 5; Park, 1983:172? 177, fi gs. 4? 6. Rhincalanus gigas Brady, 1883: Ant Rhincalanus gigas Brady, 1883:42, pl. 8, fi gs. 1? 11; Vervoort, 1951:57; 1957:34. Rhincalanus nasutus Giesbrecht, 1888: sAnt-Ntemp Rhincalanus nasutus Giesbrecht, 1888:334; 1893:152? 154, 761, pl. 3, fi g. 6; pl. 9, fi gs. 6, 14; pl. 12, fi gs. 9? 12, 14, 16, 17; pl. 35, fi gs. 46, 47, 49; Vervoort 1957:33. Scaphocalanus antarcticus Park, 1982: Ant Scaphocalanus antarcticus Park, 1982:83? 89, fi gs. 3? 7. Scaphocalanus cristatus (Giesbrecht, 1895): sAnt-Ntemp Scolecithrix cristata Giesbrecht, 1895:252? 253, pl. 2, fi gs. 6? 8; pl. 3, fi gs. 1? 5. Scaphocalanus cristatus Park, 1982:92? 95, fi g. 10. Scaphocalanus echinatus (Farran, 1905): sAnt-Ntemp Scolecithrix echinata Farran, 1905:37? 38, pl. 4, fi gs. 15? 18; pl. 5, fi gs. 12? 17. Scaphocalanus echinatus Park, 1982:101? 104, fi gs. 15? 16. Scaphocalanus elongatus Scott, 1909: Ant-Ntemp Scaphocalanus elongatus A. Scott, 1909:98, pl. 32, fi gs. 10? 16; Park, 1982:106? 108, fi gs. 18? 19. Scaphocalanus farrani Park, 1982: Ant-sAnt Scaphocalanus farrani Park, 1982:95? 101, fi gs. 11? 14. Scaphocalanus major (Scott, 1894): Ant-Trop Scolecithrix major Scott, 1894:52, pl. 3, fi gs. 24? 26; pl. 5, fi gs. 44? 45. Scaphocalanus major Park, 1982:108? 110, fi g. 20. Scaphocalanus medius (Sars, 1907): sAnt-Ntemp Amallophora media Sars, 1907:16. Scaphocalanus medius Sars, 1925:173? 174, pl. 49, fi gs. 1? 8; Park, 1982:110? 112, fi g. 21. Scaphocalanus parantarcticus Park, 1982: Ant-sAnt Scaphocalanus parantarcticus Park, 1982:89? 92, fi gs. 8? 9. Scaphocalanus subbrevicornis (Wolfenden, 1911): Ant Amallophora subbrevicornis Wolfenden, 1911:262? 263, text fi g. 37. Scaphocalanus subbrevicornis Park, 1982:117? 121, fi g. 26. Scaphocalanus vervoorti Park, 1982: Ant Scaphocalanus vervoorti Park, 1982:112? 117, fi gs. 22? 25. Scolecithricella altera (Farran, 1929): Ant-Ntemp Amallophora altera Farran, 1929:252, fi g. 19. Scolecithricella altera Park, 1980:70? 72, fi g. 22. Scolecithricella cenotelis Park, 1980: Ant Scolecithricella cenotelis Park, 1980:59? 60, fi g. 18. Scolecithricella dentata (Giesbrecht, 1893): sAnt-Ntemp Scolecithrix dentata Giesbrecht, 1893:266, pl.13, fi gs. 12, 20, 33, pl. 37, fi gs. 13? 14. Scolecithricella dentata Park, 1980:42? 43, fi g. 7. Scolecithricella dentipes Vervoort, 1951: Ant-sAnt Scolecithricella dentipes Vervoort, 1951:108, fi gs. 55? 59; Park, 1980:46? 50, fi gs. 10? 11. Scolecithricella emarginata (Farran, 1905): Ant-Ntemp Scolecithrix emarginata Farran, 1905:36, pl. 7, fi gs. 6? 17. Scolecithricella emarginata Park, 1980:61? 66, fi g. 19. Scolecithrix polaris Wolfenden, 1911:252? 253, pl. 30, fi gs. 1? 2, text fi g. 31a? e. 12_Park_pg143-180_Poles.indd 17712_Park_pg143-180_Poles.indd 177 11/17/08 8:30:33 AM11/17/08 8:30:33 AM 178 SMITHSONIAN AT THE POLES / PARK AND FERRARI Scolecithricella hadrosoma Park, 1980: Ant-Stemp Scolecithricella hadrosoma Park, 1980:66, fi g. 20. Scolecithricella minor (Brady, 1883): Ant-sArc Scolecithrix minor Brady, 1883:58, pl. 16, fi gs. 15? 16. Scolecithricella minor Park, 1980:31? 36, fi gs. 2? 3. Scolecithrix glacialis Giesbrecht, 1902:25, pl. 4, fi gs. 1? 7. Scolecithricella obtusifrons (Sars, 1905): Ant-Ntemp Amallophora obtusifrons Sars, 1905a:22. Scolecithricella obtusifrons Park, 1980:66? 70, fi g. 21. Scolecithricella ovata (Farran, 1905): Ant-Ntemp Scolecithrix ovata Farran, 1905:37, pl. 6, fi gs. 13? 18; pl. 7, fi gs. 1? 5. Scolecithricella ovata Park, 1980:58? 59, fi g. 17. Scolecithricella parafalcifer Park, 1980: Ant-Stemp Scolecithricella parafalcifer Park, 1980:50? 51, fi g. 12. Scolecithricella profunda (Giesbrecht, 1893): sAnt-Ntemp Scolecithrix profunda Giesbrecht, 1893:266, pl. 13, fi gs. 5, 26. Scolecithricella profunda Park, 1980:36? 37, fi g. 4. Scolecithricella pseudopropinqua Park, 1980: sAnt-Stemp Scolecithricella pseudopropinqua Park, 1980:51? 55, fi gs. 13? 14. Scolecithricella schizosoma Park, 1980: Ant-sAnt Scolecithricella schizosoma Park, 1980:43? 46, fi gs. 8? 9. Scolecithricella valida (Farran, 1908): Ant-sArc Scolecithrix valida Farran, 1908:55, pl. 5, fi gs. 14? 17; pl. 6, fi g. 7. Scolecithricella valida Park, 1980:55? 58, fi gs. 15? 16. Scolecithricella vervoorti Park, 1980: Ant Scolecithricella vervoorti Park, 1980:72? 74, fi gs. 23? 24. Scolecithricella vittata (Giesbrecht, 1893): sAnt-Ntemp Scolecithrix vittata Giesbrecht, 1893:266, pl. 13, fi gs. 2, 23, 32, 34; pl. 37, fi gs. 5, 8. Scolecithricella vittata Park, 1980:37? 42, fi gs. 5? 6. Scottocalanus helenae (Lubbock, 1856): sAnt-Ntemp Undina helenae Lubbock, 1856:25? 26, pl. 4, fi g. 4; pl. 7, fi gs. 1? 5. Scottocalanus helenae Park, 1983:205? 208, fi gs. 23? 24. Scottocalanus securifrons (Scott, 1894): sAnt-Ntemp Scolecithrix securifrons Scott, 1894:47? 48, fi gs. 41, 43? 47, 49? 52, 54, 56, pl. 5, fi g. 1. Scottocalanus securifrons Park, 1983:200? 205, fi gs. 19? 22. Scottocalanus thorii With, 1915: sAnt-sArc Scottocalanus thorii With, 1915:215? 219, pl. 6, fi gs. 14a? 14c; pl. 8, fi gs. 14a? 14b, text fi gs. 68? 70; Park, 1983:208? 210, fi g. 25. Spinocalanus abyssalis Giesbrecht, 1888: Ant-sArc Spinocalanus abyssalis Giesbrecht, 1888:355; 1893:209, pl. 13, fi gs. 42? 48; Damkaer, 1975:17? 20, fi gs. 4? 10, 148. Spinocalanus antarcticus Wolfenden, 1906: Ant, Arc Spinocalanus antarcticus Wolfenden, 1906:43, pl. 14, fi gs. 6? 9; Damkaer, 1975:30? 35, fi gs. 43? 68, 152; Bradford-Grieve, 1994:101, 103, fi g. 57. Spinocalanus horridus Wolfenden, 1911: Ant-Arc Spinocalanus horridus Wolfenden, 1911:216, text fi g. 7, pl. 25, fi gs. 1? 2; Damkaer, 1975:37? 41, fi gs. 69? 83, 153. Spinocalanus magnus Wolfenden, 1904: Ant-sArc Spinocalanus magnus Wolfenden, 1904:118; Damkaer, 1975: 26? 30, fi gs. 35? 42, 150. Spinocalanus terranovae Damkaer, 1975: Ant Spinocalanus terranovae Damkaer, 1975:60? 62, fi gs. 141? 147, 159. Stephos longipes Giesbrecht, 1902: Ant Stephos longipes Giesbrecht, 1902:20, pl. 2, 6? 14; Tanaka, 1960:37, pl. 14, fi gs. 1? 10. Stephos antarcticus Wolfenden, 1908: Ant Stephos antarcticus Wolfenden, 1908:24, pl. 5, fi gs. 4? 8. Subeucalanus longiceps (Matthews, 1925): sAnt-Stemp Eucalanus longiceps Matthews, 1925:127, pl. 9; Vervoort, 1957:33, fi g. 8. Subeucalanus longiceps Bradford-Grieve, 1994:88, fi gs. 40, 41, 91. Talacalanus greeni (Farran, 1905): Ant-Ntemp Xanthocalanus greeni Farran, 1905:39, pl. 8, fi gs. 1? 13; Park, 1983:325? 327, fi gs. 5? 6. Talacalanus calaminus Wolfenden, 1906, pl. 11, fi gs. 3? 5. Talacalanus greeni (Farran, 1905); Campaner, 1978:976. Temorites brevis Sars, 1900: Ant-Arc Temorites brevis Sars, 1900:100, pls. 30, 31; Vervoort, 1957:115? 118, fi gs. 102? 108. Undeuchaeta incisa Esterly, 1911: Ant-sArc Undeuchaeta incisa Esterly, 1911:319, pl. 27, fi gs. 12, 19; pl. 28, fi g. 28; pl. 29, fi g. 59; Park, 1978:183? 187, fi gs. 54? 56; Markhaseva, 1996:302, fi gs. 243? 244. Undeuchaeta major Giesbrecht, 1888: sAnt-Ntemp Undeuchaeta major Giesbrecht, 1888, p.336; Markhaseva, 1996:305, fi gs. 246? 247. Undeuchaeta plumosa (Lubbock, 1856): sAnt-Ntemp Undina plumosa Lubbock, 1856:24, pl. 9, fi gs. 3? 5; Markhaseva, 1996:310, fi gs. 248? 249. Undinella brevipes Farran, 1908: sAnt-sArc Undinella brevipes Farran, 1908:12, 50, pl. 5, fi gs. 1? 4;Vervoort, 1957:95? 96. Valdiviella brevicornis Sars, 1905: Ant-Ntemp Valdiviella brevicornis Sars, 1905a:17; 1925:101, pl. 28, fi gs. 11? 17; Park, 1978:195? 197, fi g. 61. Valdiviella insignis Farran, 1908: Ant-Ntemp Valdiviella insignis Farran, 1908:45, pl. 3, fi gs. 1? 6; Park, 1978:197? 199, fi g. 62. Valdiviella minor Wolfenden, 1911: sAnt-Ntemp Valdiviella minor Wolfenden, 1911:249, pl. 29, fi gs. 8? 11; Park, 1978:199, fi gs. 63, 64. Valdiviella oligarthra Steuer, 1904: Ant-Ntemp Valdiviella oligarthra Steuer, 1904:593, fi gs. 1? 3; Park, 1978:191? 195, fi gs. 59? 60. APPENDIX 2 This appendix lists families, alphabetically, and gen- era within families, alphabetically, with species reported from the Southern Ocean. Acartiidae Paralabidocera Aetideidae Aetideopsis Aetideus Batheuchaeta Chiridiella Chiridius Chirundina Gaetanus Euchirella Pseudeuchaeta Pseudochirella Undeuchaeta Valdiviella Arietellidae Arietellus 12_Park_pg143-180_Poles.indd 17812_Park_pg143-180_Poles.indd 178 11/17/08 8:30:33 AM11/17/08 8:30:33 AM PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 179 Augaptilidae Augaptilus Centraugaptilus Euaugaptilus Haloptilus Pachyptilus Pseudaugaptilus Bathypontiidae Temorites Calanidae Calanoides Calanus Neocalanus Candaciidae Candacia Clausocalanidae Clausocalanus Ctenocalanus Drepanopus Farrania Microcalanus Eucalanidae Eucalanus Rhincalanus Euchaetidae Paraeuchaeta Heterorhabdidae Disseta Heterorhabdus Heterostylites Paraheterorhabdus Lucicutiidae Lucicutia Megacalanidae Bathycalanus Bradycalanus Megacalanus Metridinidae Metridia Pleuromamma Nullosetigeridae Nullosetigera Phaennidae Cornucalanus Onchocalanus Talacalanus Scolecitrichidae Cephalophanes Landrumius Lophothrix Racovitzanus Scaphocalanus Scolecithricella Scottocalanus Spinocalanidae Spinocalanus Stephidae Stephos Subeucalanidae Subeucalanus Tharybidae Undinella 12_Park_pg143-180_Poles.indd 17912_Park_pg143-180_Poles.indd 179 11/17/08 8:30:34 AM11/17/08 8:30:34 AM 12_Park_pg143-180_Poles.indd 18012_Park_pg143-180_Poles.indd 180 11/17/08 8:30:34 AM11/17/08 8:30:34 AM ABSTRACT. We summarize and evaluate explanations that have been proposed to ac- count for the unusually high number of benthic marine invertebrate species in the South- ern Ocean with nonpelagic development. These explanations are divided between those involving adaptation to current conditions in this cold-water environment, selecting for nonpelagic larval development, and those involving vicariant events that either extermi- nated a high proportion of species with pelagic development (the extinction hypothesis) or enhanced speciation in taxa that already had nonpelagic development. In the latter case, glacial maxima over the Antarctic Continental Shelf in the Pliocene/Pleistocene gla- cial cycles could have created refuges where speciation occurred (the ACS hypothesis), or the powerful Antarctic Circumpolar Current passing through Drake Passage for over 30 million years could have transported species with nonpelagic development to new habitats to create new species (the ACC hypothesis). We examine the distribution and phylogenetic history of echinoderms and crustaceans in the Southern Ocean to evaluate these different explanations. We could fi nd little or no evidence that nonpelagic develop- ment is a direct adaptation to conditions in the Southern Ocean. Some evidence supports the three vicariant hypotheses, with the ACC hypothesis perhaps the best predictor of observed patterns, both the unusual number of species with nonpelagic development and the notably high biodiversity found in the Southern Ocean. INTRODUCTION The unusually high incidence of parental care displayed by marine benthic invertebrates in the Southern Ocean was fi rst noted by members of the pioneering nineteenth century expedition of the R/V Challenger (Thomson, 1876, 1885). Ex- amples were found in four of the fi ve classes of echinoderms as well as in molluscs, polychaetes, and other groups. By the end of the century, the idea was widely accepted: nonpelagic development by brooding or viviparity or within egg cap- sules was the dominant mode of reproduction by benthic marine animals, not only for Antarctic and subantarctic forms but also for cold-water species in gen- eral (Thomson, 1885; Murray, 1895; beautifully reviewed by Young, 1994). This notion was persuasively reinforced by Thorson (1936, 1950), who focused on gastropods in the Northern Hemisphere, and Mileikovsky (1971), who termed it ?Thorson?s rule.? Both Thorson (1936) and Mileikovsky (1971), however, recog- nized many exceptions, and subsequently, with more information and reanalyses John S. Pearse, Department of Ecology and Evolutionary Biology, Long Marine Labora- tory, University of California, Santa Cruz, 100 Shaffer Road, Santa Cruz, CA 95060, USA. Richard Mooi and Susanne J. Lock- hart, Department of Invertebrate Zool- ogy and Geology, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA 94118-4503, USA. Angelika Brandt, Zoologisches Institut und Zoolo- gisches Museum, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany. Correspond- ing author: J. S. Pearse (pearse@biology .ucsc.edu). Accepted 19 May 2008. Brooding and Species Diversity in the Southern Ocean: Selection for Brooders or Speciation within Brooding Clades? John S. Pearse, Richard Mooi, Susanne J. Lockhart, and Angelika Brandt 13_Pearse_pg181-196_Poles.indd 18113_Pearse_pg181-196_Poles.indd 181 11/17/08 8:37:07 AM11/17/08 8:37:07 AM 182 SMITHSONIAN AT THE POLES / PEARSE ET AL. of earlier data, the generality of Thorson?s rule weakened substantially (Pearse et al., 1991; Clarke, 1992; Hain and Arnaud, 1992; Pearse, 1994; Young, 1994; Stanwell-Smith et al., 1999; Arntz and Gili, 2001; Schluter and Rachor, 2001; Absher et al., 2003; Sewell, 2005; V?zquez et al., 2007; Fetzer and Arntz, 2008). We now know that many of the most abundant species in Antarctic waters, especially those in shallow water, have pelagic larvae as in other ar- eas of the world. Moreover, taxa in the Arctic (Dell, 1972; Fetzer and Arntz, 2008) and the deep sea (Gage and Tyler, 1991) do not have the unusually high numbers of brooding species found in the Antarctic, with the exception of peraca- rids, all of which brood and are abundant in the Arctic and deep sea, though less diverse than in the Antarctic. Indeed, as shown by Gallardo and Penchaszadeh (2001), the inci- dence of brooding species of gastropods depends at least as much on the clades present in an area as on location. Although Thorson?s rule no longer applies in general terms, it was originally based on solid observations of some unusual taxa that brood in the Southern Ocean (re- viewed by Pearse and Lockhart, 2004). Initially, the fi nd- ing of species with nonpelagic development was attributed to adaptation to conditions peculiar to polar seas (Murray, 1895; Thorson, 1936, 1950; Hardy, 1960: Pearse, 1969; Mileikovsky, 1971). However, because high incidences of brooding occur mainly in Antarctic waters and not in the Arctic (Ludwig, 1904; ?stergren, 1912; Dell, 1972), it be- came clear that something besides adaptation to ?harsh? polar conditions had to be involved. Thorson (1936), rec- ognizing the difference between the two polar seas, sug- gested that the Arctic fauna, being younger than those around the Antarctic, had not had as much time to adapt; this explanation was accepted by others (e.g., Arnaud, 1974; Picken, 1980). Nevertheless, as recognized by Dell (1972), the discrepancy between the two polar seas meant that the unusual incidence of nonpelagic development in the Southern Ocean was not likely to be the consequence of simple adaptation to some general polar conditions. While there can be little doubt that developmental mode is infl uenced by, and at least initially determined by, natural selection, the adaptive nature of one particu- lar mode over another has been subject to considerable speculation and debate (Strathmann, 1993; Havenhand, 1995; Wray, 1995; Gillespie and McClintock, 2007). Pelagic development, either with feeding or nonfeeding larvae, has usually been assumed to be plesiomorphic, and benthic development has been assumed to be derived (J?gersten, 1972; Villinski et al., 2002; Gillespie and Mc- Clintock, 2007). Moreover, once lost, planktotrophic de- velopment is rarely reacquired (Strathmann, 1978; Reid, 1990; Levin and Bridges, 1995; but see Collin et al., 2007), and this generalization probably applies to pelagic development in general. Consequently, the occurrence of benthic development in a taxon may be an adaptation to particular conditions (e.g., oligotrophic water or offshore currents), or it may be a phyletic constraint refl ecting earlier adaptations that no longer apply. Paleontological evidence suggests that species of marine molluscs with nonpelagic development had smaller distributions and were more susceptible to extinction than those with pe- lagic development (Jablonski and Lutz, 1983; Jablonski and Roy, 2003); presumably, these had more genetically fragmented populations as well. An alternative explanation to the unusually numerous brooding species in the Southern Ocean is that their high numbers are the consequence of populations being repeat- edly fragmented, with isolated units forming new species. That is, nonpelagic development in the Southern Ocean might not refl ect adaptation scattered among several clades, as it does elsewhere (e.g., Byrne et al., 2003; Col- lin, 2003), but rather, it may occur mainly in relatively few clades in which species proliferated. Moreover, some of these species-rich, brooding clades could contribute sub- stantially to the unexpected high species diversity found in the Southern Ocean (Brandt et al., 2007a, 2007b; Rogers, 2007). Indeed, in some taxa, species-rich clades of brood- ers constitute most of the species (e.g., echinoids: Poulin and F?ral, 1996; David et al., 2003, 2005; crustaceans: Brandt, 2000; Brandt et al., 2007a, 2007b). Consequently, the occurrence of many species with nonpelagic develop- ment may not be due to specifi c adaptations to conditions in the Antarctic but, instead, may be a consequence of iso- lation after vicariant events that now or in the past led to their proliferation. In this paper we evaluate and compare several adaptation versus vicariant explanations for the occurrence of species-rich clades in the high latitudes of the Southern Ocean. PROPOSED EXPLANATIONS ADAPTATION Although some aspect of the current polar environ- ment has usually been assumed to have led to the selection of nonpelagic development in the Southern Ocean, iden- tifi cation of the responsible agents has been elusive. The problem is compounded because unusually high numbers of brooding species are found in Antarctic and subantarc- tic waters but not in either the Arctic or deep sea, the other areas of the world ocean with cold water year-round. We 13_Pearse_pg181-196_Poles.indd 18213_Pearse_pg181-196_Poles.indd 182 11/17/08 8:37:07 AM11/17/08 8:37:07 AM BROODING AND SPECIES DIVERSITY IN THE SOUTHERN OCEAN 183 briefl y consider below some of the ideas that have been proposed, including those that apply to cold-water envi- ronments in general. Low Temperature Murray (1895:1459) suggested simply that ?animals with pelagic larvae would be killed out or be forced to migrate towards the warmer tropics? when temperatures cooled, to be replaced by animals without larvae existing below the ?mud-line? where he thought very few animals had pelagic larvae. Similarly, Thatje et al. (2005b) argue that the predominance of developmental lecithotrophy in the Antarctic is the consequence of the near-complete ex- tinction of benthic communities during glacial maxima and recolonization from deeper waters where species had un- dergone an ?evolutionary temperature adaptation? that led to lecithotrophy. However, no evidence supports the idea that either nonpelagic development or lecithotrophy is an adaptation to low temperature, and the fact that a wide variety of both planktotrophic and lecithotrophic pelagic larvae have been found in both Antarctic and Arctic waters (e.g., Thorson, 1936; Stanwell-Smith et al., 1999; Sewell, 2005; Palma et al., 2007; V?zquez, 2007; Fetzer and Arntz, 2008) persuasively indicates that marine invertebrate lar- vae are able to survive and grow at freezing temperatures? even under high pressures found in the deep sea (Tyler et al., 2000), where many species have pelagic, planktotro- phic larvae (Gage and Tyler, 1991; however, see below). Low Temperature and Slow Development Many studies have shown that larval development is greatly slowed at very low temperatures (e.g., Hoegh- Guldberg and Pearse, 1995; Peck et al., 2007), and the metabolic basis of this effect is gradually being sorted out (e.g., Peck, 2002; Clarke, 2003; Peck et al. 2006; Pace and Manahan, 2007). The longer larvae are in the plankton, the greater the chance that they will perish by predation or be swept away from suitable settling sites. Indeed, Smith et al. (2007) argued that lecithotrophic development might be selected because eliminating the feeding stage substan- tially shortens the time larvae spend in the plankton, a particular advantage for polar areas where development is slow. Going one step further, nonpelagic development eliminates loss in the plankton altogether. However, not only would this explanation apply to the cold-water envi- ronment of the Arctic and deep sea as well as to the Antarc- tic, but as mentioned above, many particularly abundant polar species do have slow-developing planktotrophic pe- lagic larvae; long periods of feeding in the plankton do not necessarily appear to be selected against. Low Temperature, Slow Development, and Limited Larval Food Thorson (1936, 1950) developed the idea that plank- totrophic larvae would be food limited in polar seas be- cause phytoplanktonic food is available only during the summer plankton bloom, too briefl y for such larvae to complete their slow development. This durable hypoth- esis remains current (e.g., Arntz and Gili, 2001; Thatje et al., 2003, 2005b), although little or no evidence supports it. Indeed, planktotrophic larvae of a wide range of taxa are well known in polar seas (see above). Moreover, ex- tremely low metabolic rates of gastropod and echinoid lar- vae, indicative of very low food requirements, have been demonstrated by Peck et al. (2006) and Pace and Mana- han (2007), respectively. There is no evidence that other planktotrophic larvae of polar seas are food limited either. In addition, this proposal is not specifi c to the Southern Ocean. Finally, it applies only to planktotrophic larval development, not lecithotrophic pelagic development; our concern here is pelagic and nonpelagic development, not mode of nutrition for developing embryos or larvae. Low Adult Food Supply Chia (1974) suggested that poor nutritional conditions for adults might favor nonpelagic development on the as- sumption that adults require more energy to produce the multitude of pelagic larvae needed to overcome high larval mortality in the plankton than to produce a few protected offspring. Such conditions do prevail in polar seas, where primary production is extremely seasonal (Clarke, 1988), or especially during periods of maximal multiyear sea ice and glacial expansion during the Pliocene/Pleistocene ice ages (see below). However, studies on a poecilogonous species of polychaete indicate that nutritional investment is higher in the form that produces lecithotrophic larvae than in one that produces planktotrophic larvae (Bridges, 1993), coun- tering Chia?s (1974) assumption. Moreover, even if true, the argument applies to both polar seas and is not specifi c to the Southern Ocean. Finally, there is little evidence that polar species are food limited over an entire year. Large Egg Sizes It has long been known that egg size and, presum- ably, energy investment into individual eggs increase with 13_Pearse_pg181-196_Poles.indd 18313_Pearse_pg181-196_Poles.indd 183 11/17/08 8:37:08 AM11/17/08 8:37:08 AM 184 SMITHSONIAN AT THE POLES / PEARSE ET AL. increasing latitude (reviewed by Laptikhovsky, 2006). If more energy is allocated to each egg, fecundity is low- ered. Moreover, larger eggs require more time to com- plete the nonfeeding phase of development than smaller eggs (Marshall and Bolton, 2007), increasing the risk of embryonic/larval mortality while in the plankton. With lower fecundity and increased risk of mortality, there could be strong selection for nonpelagic development, eliminating mortality in the plankton altogether. While the underlying reason why egg size increases with lati- tude remains to be understood, it could be a factor lead- ing to nonpelagic development. However, this explana- tion also applies to both polar regions and not solely to the Southern Ocean. Small Adult Size It is also well known that taxa composed of smaller in- dividuals tend to have nonpelagic development, while those with larger individuals tend to have planktotrophic, pelagic development (reviewed by Strathmann and Strathmann, 1982). This observation is also based on fecundity: small animals cannot produce enough offspring for any of them to have much chance of surviving the high mortality faced in the plankton. However, there are many examples of spe- cies comprised of very small individuals producing plank- totrophic larvae, making generalization diffi cult. Neverthe- less, most species in some major taxa (e.g., bivalves: Clarke, 1992, 1993) in the Southern Ocean are composed of very small individuals, so this explanation could apply to them, at least in terms of factors originally selecting for nonpe- lagic development. Low Salinity ?stergren (1912) suggested that melting ice during the summer would create a freshwater layer unfavorable to pelagic larvae and therefore could be a factor select- ing against them. Thorson (1936), Hardy (1960), Pearse (1969), and Picken (1980) also considered low salinity to be a factor selecting against pelagic development in polar seas. The large rivers fl owing into the Arctic Ocean should make low salinity an even greater problem there than around the Antarctic. Yet, as with most of the adap- tationist explanations focusing on polar conditions, the fact that nonpelagic development is less prevalent in the Arctic than in the Antarctic undermines this explanation. Thus, low salinity is not likely to be an important fac- tor selecting for nonpelagic development in the Southern Ocean. Narrow Shelf Recognizing that the presence of unusually high num- bers of species with nonpelagic development is mainly a feature of the Antarctic, especially the subantarctic, rather than the Arctic, ?stergren (1912) also proposed that the narrow shelf and the strong winds blowing off the conti- nent would drive larvae offshore, away from suitable set- tling sites. Consequently, there would be strong selection against pelagic larvae in the Antarctic but not in the Arc- tic. This was the fi rst attempt to explain the high number of brooding species specifi cally for the Southern Ocean. However, the idea was quickly discounted by Mortensen (1913), who pointed out that it should apply as well to remote oceanic islands in the tropics, where pelagic devel- opment was already well known. SELECTIVE EXTINCTION Another possibility is that events in the past led to the extinction of many or most species with pelagic develop- ment in the Antarctic, leaving a disproportionate number of species with nonpelagic development. This proposal by Poulin and F?ral (1994) argues that pelagic development was not adaptive, while nonpelagic development was neu- tral at certain times in the past in the Southern Ocean. It was developed further by Poulin and F?ral (1996), Pou- lin et al. (2002), and Thatje et al. (2005b) and is based on the fi nding that during the glacial maxima of the late Quaternary ice ages, grounded ice extended to the edge of the Antarctic shelf (Clarke and Crame, 1989), obliterating most life on the shelf. During such times, thick, multiyear sea ice probably occurred year-round and extended far into the Southern Ocean surrounding the Antarctic, block- ing sunlight and photosynthesis beneath it. Consequently, there would be little (only laterally advected) phytoplank- tonic food to support planktotrophic larvae, and selection would be strong for lecithotrophic development, whether pelagic or benthic. Of course, primary production is nec- essary to support populations of juveniles and adults as well, and although massive extinction would be expected during the glacial maxima of the Pliocene/Pleistocene re- gardless of developmental mode, this apparently did not happen (Clarke, 1993). Poulin and F?ral (1994, 1996) suggested that selective extinction of species with planktotrophic larvae during the glacial maxima would leave behind species with non- pelagic development, but as Pearse and Lockhart (2004) pointed out, such selective extinction would leave species with both pelagic and nonpelagic lecithotrophic develop- 13_Pearse_pg181-196_Poles.indd 18413_Pearse_pg181-196_Poles.indd 184 11/17/08 8:37:08 AM11/17/08 8:37:08 AM BROODING AND SPECIES DIVERSITY IN THE SOUTHERN OCEAN 185 ment. Consequently, the selective extinction hypothesis is inadequate to explain the unusual abundance of spe- cies with nonpelagic development. However, in the case of echinoids (the taxon of concern for Poulin and F?ral, 1994, 1996), there are very few species anywhere with lec- ithotrophic pelagic larvae? and none in the Antarctic? so the selective extinction hypothesis applies at least partly to echinoids. Similarly, the high diversity of peracarid crusta- ceans, most of which brood, and the paucity of decapod crustaceans, most of which have planktotrophic larvae, also may be the consequence, at least in part, of selective extinction (Thatje et al., 2003, 2005b). Recognizing that brooding might have originated outside the Antarctic, Dell (1972), Arnaud (1974), and Picken (1980) suggested that brooding species could have colonized the Southern Ocean from elsewhere after mas- sive extinctions, perhaps by rafting (see Thiel and Gutow, 2005). On the other hand, polar emergence from the deep sea following the retreat of multiyear sea ice in intergla- cial periods might have taken place for some taxa, which subsequently speciated on the Antarctic shelf (e.g., isopod families Munnopsidae, Desmosomatidae, Ischnomesidae, e.g., Br?keland, 2004; Raupach et al., 2007). ENHANCED SPECIATION In contrast to enhanced extinction of species with pelagic development, which would leave behind a dispro- portionate number of species with nonpelagic develop- ment, speciation could be enhanced by conditions in the Southern Ocean, in the past or persisting to the present, to produce species-rich clades of taxa with nonpelagic devel- opment. Nonpelagic development could have developed well before the Southern Ocean cooled, or even elsewhere altogether, but spread via a founding species to the South- ern Ocean and then undergone radiation. Regardless of where or how nonpelagic development originated, if this is the case, we have not only an explanation of the un- usually high number of species with nonpelagic develop- ment but also, perhaps, an explanation for the unexpected high species diversity in the Southern Ocean (Brandt et al., 2007a, 2007b; Rogers, 2007). At least two different scenarios about how this might occur are specifi c to the Southern Ocean. Isolation and Speciation on the Antarctic Continental Shelf (the ACS Hypothesis) Clarke and Crame (1989, 1992, 1997), Brandt (1991, 2000), and Thatje et al. (2005b) pointed out that during the glacial maxima, grounded ice probably did not com- pletely cover the shelf areas around the Antarctic conti- nent. Instead, some isolated areas would likely be open and habitable under the ice, as seen today under ice shelves (e.g., Littlepage and Pearse, 1962; Post et al., 2007). These areas could behave as ?islands? with remnants of the pre- viously more widespread shelf fauna. Species with non- pelagic development would be effectively isolated. With the retreat of the grounded glaciers, the shelf fauna would reconnect, mixing the newly formed species as they ex- panded around the continent. Similar phenomena might be happening now after the disintegration of the Larsen A and B ice shelves in 2002. During the height of interglacial periods, when there was a minimum of ice cover, the Wed- dell and Ross seas could have been connected (Scherer et al., 1998; Thomson, 2004), further mixing species, which would be fragmented again during the subsequent gla- cial cycle. Clarke and Crame (1989) proposed that such repeated cycles of glacial advances and retreats over the shelf could favor speciation, and Clarke and Crame (1992, 1997) further developed this idea and suggested that such oscillation would act as a ?species diversity pump.? It would be most effective, however, for species having lim- ited dispersal capabilities, such as those with nonpelagic development. Isolation during glacial maxima is not the only pos- sibility for fragmenting populations on the Antarctic Con- tinental Shelf. At present, most shallow-water habitats (H11021150 m) around the Antarctic continent are covered by grounded ice or fl oating ice shelves, and only scattered fragments of suitable habitats remain. Ragu?-Gil et al. (2004) found that three such habitats, one on the west side of the Antarctic Peninsula and two others on the eastern coast of the Weddell Sea, support very different faunas. The biotas in the two habitats in the Weddell Sea differ as much from each other as they do from the one on the Antarctic Peninsula. According to Ragu?-Gil et al. (2004), these differences indicate limited exchange due at least in part to a predominance of species with nonpelagic larvae. Similar differences were detected for isopod composition at sites around the Antarctic Peninsula and in the Weddell Sea (Brandt et al., 2007c). Such isolation could lead to speciation, particularly of cryptic species formed by non- selective processes (e.g., genetic drift). Speciation of fragmented populations on the Antarctic Continental Shelf, the ACS hypothesis, would result in an increase of shelf species, so that the greatest species rich- ness would be expected on the shelf, with decreasing rich- ness down the slope into deeper depths. That was found to be the case for amphipods (Brandt, 2000), polychaetes 13_Pearse_pg181-196_Poles.indd 18513_Pearse_pg181-196_Poles.indd 185 11/17/08 8:37:08 AM11/17/08 8:37:08 AM 186 SMITHSONIAN AT THE POLES / PEARSE ET AL. (but not isopods or bivalves) (Ellingsen et al., 2007), and many other taxa (Brandt et al., 2007a) sampled in the At- lantic sector of the Southern Ocean. In addition, because most of the glacial cycles occurred during the Pliocene and Pleistocene over only the past few million years, ge- netic divergence of these fragmented populations would be relatively slight, and very similar or cryptic sister spe- cies would be predicted. Molecular analyses have revealed cryptic species in isopods, which brood (Held, 2003; Held and W?gele, 2005), and a bivalve that broods (Linse et al., 2007) as well as in a crinoid, which has pelagic, lecithotro- phic larvae (Wilson et al., 2007). Isolation and Speciation via the Antarctic Circumpolar Current (the ACC Hypothesis) Pearse and Bosch (1994) analyzed available data for mode of development in shallow-water Antarctic and sub- antarctic echinoderms (128 species) and found the highest proportion of species with nonpelagic development in the region of the Scotia Arc (65%), not the Antarctic continent or subantarctic islands (42% each). This pattern led them to focus on Drake Passage and the powerful Antarctic Cir- cumpolar Current (ACC) that has been fl owing through it for more than 30 million years (Thomson, 2004). They proposed that individuals of species with nonpelagic devel- opment could be rafted infrequently to other downstream habitats and could become established to form new iso- lated populations, i.e., new species. Moreover, tectonic ac- tivity in the Scotia Arc region has continually formed new habitats as crustal plates shifted, which also infl uenced complex eddies as water fl owed through Drake Passage (Thomson, 2004). With more than 30 million years since the ACC broke through Drake Passage, many new species could form and accumulate. Pearse and Lockhart (2004) reviewed these ideas, found further support for them, and suggested ways to test them using cidaroids. The ACC hypothesis predicts that species richness would consist of species-rich clades of taxa with nonpe- lagic development and would not be an accumulation of many species-poor clades with a variety of reproductive modes, including nonpelagic development, which ap- pears to be the case (see below). Moreover, species rich- ness would be greatest within and east of the Scotia Arc, downstream from Drake Passage. The Scotia Arc region, in fact, appears to be unusually diverse (Barnes, 2005; Barnes et al., 2006; Linse et al., 2007). Conversely, species diversity should be lower upstream, on the western side of the Antarctic Peninsula, and that is exactly the pattern Ragu?-Gil et al. (2004) found in their analysis of three shallow-water communities. Similar differences in species richness between the eastern Weddell Sea and the western coast of the Antarctic Peninsula were reported by Star- mans and Gutt (2002). On a different scale, Linse et al. (2006) likewise found the highest diversity of molluscs to be in the Weddell Sea, east of the Scotia Arc, and the lowest on the western side of the Antarctic Peninsula (al- though this might have been due to sampling discrepan- cies). It can also be predicted that because the ACC hits the Antarctic Peninsula as it fl ows around the continent, it could carry species to the western side of the peninsula, where they might accumulate (A. Mahon, Auburn Univer- sity, personal communication). In addition, because the ACC is funneled through the whole of Drake Passage, the ACC hypothesis does not necessarily predict a depth gradient of species richness, in contrast to the ACS hypothesis. No depth gradient is seen for isopods and bivalves (Ellingsen et al., 2007). Indeed, the ACC hypothesis may account for the unexpected high species diversity recently documented for some deep-sea taxa in the Atlantic portion of the Southern Ocean (Brandt et al., 2007a, 2007b). Finally, because the ACC continues to this day, it would not be unexpected for species to have formed, as described above, at any time over the past 30 million years, including within the past few million years, so that closely related cryptic species would be found as well as more dis- tantly related species, all in the same clade. Unlike the ACS hypothesis, the ACC hypothesis predicts the existence of a spectrum of variously diverged species within the clades. That result is what has been found in Lockhart?s (2006) analysis of brooding cidaroids, the fi rst thorough phylo- genetic analysis of a major clade of brooders within the Southern Ocean (see below). Species with nonpelagic development are thought to be prone to high extinction rates because they typically have small population sizes and limited distributions, which make them particularly susceptible to environmen- tal change (Jablonski and Lutz, 1983). Poulin and F?ral (1994) suggested that because of such susceptibility, any enhanced speciation rate in the Southern Ocean would be countered by a high extinction rate. Consequently, they rejected an enhanced speciation model for explaining high species diversity in clades with nonpelagic development. However, with the ACC in effect for over 30 million years, the Southern Ocean has been an extraordinarily stable marine environment. Jeffery et al. (2003) proposed an idea similar to the ACC hypothesis to explain the high propor- tion of brooding early Cenozoic echinoids that occurred on the southern coast of Australia after Australia sepa- 13_Pearse_pg181-196_Poles.indd 18613_Pearse_pg181-196_Poles.indd 186 11/17/08 8:37:09 AM11/17/08 8:37:09 AM BROODING AND SPECIES DIVERSITY IN THE SOUTHERN OCEAN 187 rated from the Antarctic. Strong currents swept through the Tasmanian Gateway then and could have swept in- dividuals with nonpelagic development to new habitats, where they would have potentially formed new species. McNamara (1994) earlier recognized the importance of the stability provided by the strong, constant current through the Tasmanian Gateway for favoring the accumu- lation of brooding echinoids; he suggested that their later disappearance was a result of the widening of the gateway and a decrease in the environmental stability. Similarly, we suggest that the ACC fl owing through Drake Passage provides conditions both for enhancing speciation and for tempering extinction. EVALUATING THE EXPLANATIONS The proposed explanations above for the unusual abundance of species with nonpelagic development in the Southern Ocean are not mutually exclusive of each other, and one or more may apply to one or more taxa. However, with recent advances in molecular phylogenetic analyses (Rogers, 2007), these proposed explanations may be bet- ter evaluated than was possible earlier. For example, (1) if nonpelagic development is scattered within taxa found in widely distributed clades and these taxa are found both within and outside the Southern Ocean, such a mode of development is not likely to be an adaptation to condi- tions in the Southern Ocean. (2) If taxa with nonpelagic development in widely distributed clades are restricted to both polar environments and the deep sea, nonpelagic de- velopment might be an adaptation to cold water; if they are only in the Southern Ocean, specifi c conditions around the Antarctic would more likely be involved. (3) If nonpe- lagic development is found in all the taxa of clades found in both the Southern Ocean and elsewhere, where the basal taxa are found may indicate where the trait origi- nated, and conditions there might be involved in the selec- tion of nonpelagic development. (4) If nonpelagic develop- ment is found disproportionately more in Southern Ocean taxa of clades than elsewhere, either this development is a consequence of adaptation to conditions specifi c to the Southern Ocean, or it is the result of extinction of taxa with pelagic development. (5) If nonpelagic development is found in many taxa of clades in the Southern Ocean but only in a few taxa of basal clades found elsewhere, the Southern Ocean taxa may have proliferated because of unusual conditions there (not necessarily because non- pelagic development was adaptive). (6) If most taxa with nonpelagic development appeared only over the past few million years, when massive glacial advances and retreats occurred, they may have been generated on the Antarctic Continental Shelf when the glacial advances separated and fragmented populations (the ACS hypothesis). (7) If the taxa appeared more or less steadily since Antarctica sepa- rated from South America, about 30 million years ago, and are most abundant in and east of the Scotia Arc, they may have been generated by infrequently rafting with the ACC to new locations (the ACC hypothesis). SELECTED TAXA Below we review some of the information now avail- able for taxa of two major groups in the Southern Ocean: echinoderms and crustaceans. Species in these taxa are major components of the Southern Ocean biota, and they are relatively well known. Moreover, phylogenetic analy- ses are now available for some groups within them, in- cluding speciose, brooding clades. Other taxa could also be evaluated for a stronger comparative analysis, in par- ticular, molluscs, pycnogonids, and teleosts; we hope that research is done by others. ECHINODERMS Nonpelagic development in echinoderms caught the attention of naturalists with the Challenger expedition in the nineteenth century (Thomson, 1876, 1885; Murray, 1895), setting the foundation for what became ?Thorson?s rule.? Echinoderms now are among the fi rst groups of ani- mals in the Antarctic to have their phylogenetic relation- ships documented. Echinoids, in particular, are revealing. Only four major clades are present in the Southern Ocean, echinids, cidaroids, holasteroids, and schizasterids (Da- vid et al., 2003, 2005). The near absence of other clades suggests either that major extinctions have occurred or that other taxa did not fi nd a foothold in the Southern Ocean. It is interesting to note that there are presently no clypeasteroids (sand dollars and allies) in Antarctica to- day, in spite of their ubiquity in cold waters both in the past and present, and that at least one species has been recorded from the Paleogene of Black Island, McMurdo Sound (Hotchkiss and Fell, 1972). Hotchkiss (1982) used this and other fossil evidence to call into question the sup- posed slow rate of evolution in cidaroids and any connec- tion between the fossil Eocene faunas of Australasia and those of the so-called ?Weddellian Province? of the South- ern Ocean. Hotchkiss (1982:682) pointed out that any supposed ?shallow-marine connection had disappeared 13_Pearse_pg181-196_Poles.indd 18713_Pearse_pg181-196_Poles.indd 187 11/17/08 8:37:09 AM11/17/08 8:37:09 AM 188 SMITHSONIAN AT THE POLES / PEARSE ET AL. by middle Oligocene time because there is evidence for the deep-fl owing Antarctic Circumpolar Current south of the South Tasman Rise at that time.? Of the three major clades, there are only seven pres- ently recognized species of echinids, all in the genus Ster- echinus. Phylogenetic analysis using mitochondrial DNA sequences indicates that the genus diverged from Lox- echinus in South America 24? 35 million years ago, when the Antarctic separated from South America (Lee et al., 2004). Two species, Sterechinus neumayeri and S. antarct- icus, are abundant and widespread around the continental shelf (Brey and Gutt, 1991). The former is known to have typical echinoid planktotrophic development (Bosch et al., 1987). The other species are less well known and are taxonomically questionable but almost certainly also have pelagic development. Although an extensive revision is pending (see Lock- hart, 2006), as of 2005, there were more than 20 recog- nized cidaroid taxa, and the vast majority of those have been recorded to be brooders? and present evidence (Lock- hart, 2006) strongly suggests that all of them are brooders. A recent phylogenetic cladogram developed by Lockhart (2006) used fossils and a penalized likelihood analysis of CO 1 , Cytochrome b (Cytb), and 18-s mitochondrial se- quences to establish divergence times for the taxa of cida- rids (see Smith et al., 2006, for an evaluation of using fos- sils for dating cladograms). The dated cladogram revealed that Southern Ocean cidaroids are monophyletic with the most likely sister taxon being the subfamily Goniocidari- nae, now found in the southwest Pacifi c, including New Zealand and Australia, but not in the Southern Ocean. A few species of goniocidarines are known to brood, but it is not yet known whether these are sister species to the Southern Ocean clade (making goniocidarines paraphy- letic). The oldest fossil goniocidarine, from the Perth Ba- sin of Western Australia when Australia and Antarctica were connected, is more than 65 million years old, and the oldest cidaroid in the Southern Ocean clade is Austro- cidaris seymourensis, from Eocene deposits on Seymour Island in the Scotia Arc, dated at 51 million years ago. Austrocidaris seymourensis had distinctive aboral brood chambers, showing that brooding was established in this clade long before cooling began. Consequently, brooding in these animals is not an adaptation to present-day condi- tions in the Southern Ocean. Lockhart (2006) also showed that there are two sister clades of Southern Ocean cidaroids: the subfamily Astro- cidarinae with two to three recently diverged species in a single genus (Austrocidaris) found in subantarctic waters on the northern edge of the Scotia Arc and the sub family Ctenocidarinae with more than 20 species in at least fi ve genera found in the southern and eastern portions of the Scotia Arc and around the Antarctic Continent. More- over, clades within the ctenocidarines diverged more or less steadily over the last 30 million years, that is, since the Antarctic Circumpolar Current was established. This pat- tern is precisely what the ACC hypothesis predicts. Among the 16 species of Holasteroida found south of the convergence, very few are found at depths less than 2000 m. Only three genera occur in relatively shallow wa- ters: Pourtalesia, Plexechinus, and two of the three known species of Antrechinus. With the exception of the latter two genera, all holasteroids belong to widespread deep- sea clades that occur well north of the Southern Ocean, and none are known to have nonpelagic, lecithotrophic development. However, within Antrechinus, we fi nd the most extreme form of brooding known in the Echinoi- dea? species that brood their young internally and ?give birth? (David and Mooi, 1990; Mooi and David, 1993). The two species known to brood are found no deeper than 1500 m. The third species ascribed to Antrechinus, A. dry- galskii, was only provisionally considered a plesiomorphic sister group to these remarkable brooders (Mooi and Da- vid, 1996) and occurs below 3000 m. There are no known fossil holasteroids from the Antarctic region. There are 30 recognized species of schizasterid Spatan- goida recorded by David et al. (2003, 2005) to occur in the Antarctic region. These are distributed in seven genera: Abatus with 11 species, Amphipneustes with 9, Tripylus with 4, Brisaster with 2, and Brachysternaster, Delopa- tagus, Genicopatagus, and Tripylaster each with a single species. Most phylogenetic analyses recognize Brisaster and Tripylaster as a monophyletic assemblage that is, at best, a sister taxon to the rest of the Antarctic Schizasteridae (F?ral, et al., 1994; Hood and Mooi, 1998; Madon-Senez, 1998; David et al., 2005; Stockley et al., 2005). The ranges of Brisaster and Tripylaster are best considered subantarctic, as there is only a single record from south of 55?S, and none have been recorded in the shelf regions of the Antarctic con- tinent. Interestingly, these genera are the only species not known to brood. All other schizasterids have nonpelagic development, brooding the young in well-developed mar- supia in the aboral, ambulacral petaloid areas (Magniez, 1980; Schatt, 1988; Pearse and McClintock, 1990; Poulin and F?ral, 1994; David et al., 2005; Galley et al., 2005). The brooding schizasterids almost undoubtedly con- stitute a single clade (Madon-Senez, 1998; David et al., 2005). Recognizing early on the need for understanding evolutionary history to understand their phylogenies, F?ral et al. (1994) compared RNA sequences in species of 13_Pearse_pg181-196_Poles.indd 18813_Pearse_pg181-196_Poles.indd 188 11/17/08 8:37:10 AM11/17/08 8:37:10 AM BROODING AND SPECIES DIVERSITY IN THE SOUTHERN OCEAN 189 the four main genera of brooding schizasterids then recog- nized in the Southern Ocean, supporting the monophyly of the brooding genera but partially undermining the monophyly of some of the constituent genera and there- fore reinforcing the later morphological work of Madon- Senez (1998). Fossils assigned to Abatus, with distinctive brood chambers, are known from the Eocene of Seymour Island on the Scotia Arc (McKinney et al., 1988). Consequently, as with the cidaroids, brooding appeared in these animals well before the Southern Seas cooled, and that mode of development cannot be attributed to polar conditions. Poulin and F?ral (1994) showed that populations of the brooding schizasterid echinoid Abatus cordatus in embay- ments around Kerguelen Island are genetically distinct, pre- sumably because of limited gene fl ow. Consequently, there is genetic differentiation in these populations of brooding echinoids, and it is likely to be occurring with other brood- ing species with limited dispersal elsewhere in the South- ern Ocean, leading to many shallow divergences in genetic structure. This is also borne out by morphological variation among specimens from different regions (Madon-Senez, 1998) and the small amounts of morphological divergence among the species themselves (David et al., 2005). Among the brooding schizasterids, very few have ranges west of the Antarctic Peninsula. Most are distrib- uted east of Drake Passage, along the South Shetlands and eastward through the Weddell region. For example, the genus Amphipneustes does not seem to occur immediately to the west of the peninsula but has abundant representa- tion to the east of the Drake Passage, with major centers of diversity in the South Shetlands and in the region of the Weddell Sea (Figure 1). This pattern is repeated for Aba- tus and the other brooding schizasterid genera. Although sampling bias could remain a mitigating factor in the ac- curacy of these distributions, we do not believe that is the case for echinoids because David et al. (2003) shows that forms such as the echinids are well represented to the west of the peninsula. Even the most diffi cult taxa to sample, the abyssal holasteroids, are almost evenly distributed around Antarctica, with no obvious gaps in the overall distribution of this clade. The implication is that the ranges of these brooding forms are being infl uenced by the prevailing ACC, which tends to force the ranges ?downstream? of the Drake Pas- sage. The precise mechanism by which brooding schizaste- rids are redistributed and then speciate remains unknown, but it does not overextend present data to suggest that once established, new populations of brooding forms can rapidly diverge from the originating population. In addition to echinids, cidaroids, holasteroids, and schizasterids, there are a few other echinoid taxa known from the deeper portions of the Southern Ocean. One species of echinothurioid is known (Mooi et al., 2004); all species so far studied in this monophyletic, wide- spread, mostly deep-water clade have pelagic, lecithotro- phic development, and the Antarctic species presumably does as well. Unusual brooding in the Southern Ocean was also highlighted in the Challenger expedition reports for the other classes of echinoderms (Thomson, 1885). However, to date, there has been no phylogenetic analysis examining whether brooders belong to a few speciose clades in these classes, as is becoming evident for echinoids. In addition, members of these other classes do not have as good a fossil record, and they do not have fossilizable structures indica- tive of brooding, as do many echinoids. Nevertheless, if the major taxonomic groups of these classes are monophyletic, the brooding species do belong to a few speciose clades. The majority of Southern Ocean asteroids, for ex- ample, are forcipulates in the family Asteriidae. Brood- ing is rare in asteriids in most of the world, limited to the speciose genus Leptasterias in north temperate/polar wa- ters (Foltz et al., 2008) and several species in genera that are mainly Antarctic/subantarctic but are also found in southern South America (e.g., Anasterias, Gil and Zaixso, FIGURE 1. Distribution of nine species of Amphipneustes around Antarctica. Data were compiled from David et al. (2003) and Polarstern and Antarctic Marine Living Resources expeditions. 13_Pearse_pg181-196_Poles.indd 18913_Pearse_pg181-196_Poles.indd 189 11/17/08 8:37:10 AM11/17/08 8:37:10 AM 190 SMITHSONIAN AT THE POLES / PEARSE ET AL. 2007; Diplasterias, Kim and Thurber, 2007) and southern Australia (Smilasterias, Rowe and Gates, 1995; Komatsu et al, 2006). However, most if not all species of asteriids in the Southern Ocean are brooders. Arnaud (1974) lists 22 species, Pearse and Bosch (1994) list 25 species, and Clarke and Johnston (2003) report that there are approxi- mately 37 species of asteriids in the Southern Ocean. Foltz et al. (2007) analyzed 31 species of forcipulates using mitochondrial and nuclear sequences and found that 30 formed a clade. Only three were Southern Ocean species (Psalidaster mordax, Cryptasterias turqueti, and Notaste- rias pedicellaris), but they formed a clade within the for- cipulate clade. There are 11 brooding species of asteroids on the subantarctic islands; seven of them are in the as- teriid genus Anasterias (Pearse and Bosch, 1994). Of the 24 species of brooding asteroids known from Antarctic waters, 18 are asteriids, and 13 of these are in two genera, Diplasterias and Lyasasterias (Pearse and Bosch, 1994). Moreover, 19 of the 24 brooding asteroids in Antarctic waters are found in the Scotia Arc region, as would be expected from the ACC hypothesis. On the other hand, most of the genera with brooding species are circumpolar (Clark, 1962; C. Mah, Smithsonian Institution, personal communication), and it may be too early to conclude that there is a disproportional number of species in the Scotia Arc region, which has been most heavily sampled to date. There is evidence that even brooding species of aster- oids are capable of wide dispersal. Diplasterias brucei, for example, is not only found around the Antarctic conti- nent and in the southern portion of the Scotia Arc but also north of the polar front on Burdwood Bank and in the Falklands Island (Kim and Thurber, 2007). Such a wide distribution by a brooding species suggests unusual capa- bilities of dispersal, such as by rafting. Moreover, genetic analyses would be expected to show considerable genetic differentiation, as found for Abatus cordatus at Kerguelen Islands (Poulin and F?ral, 1994). Such analyses would be most welcome. Although some of the asteriid species with nonpelagic development are commonly found in the Southern Ocean, the most frequently encountered asteroids on the Antarc- tic shelf are species of Odontasteridae, especially Odon- taster validus, which like the echinid echinoid Sterechinus neumayeri, is found around the Antarctic continent, often in very high numbers. There are about 11 species of odon- tasterids in the Southern Ocean (Clarke and Johnston, 2003), two or three in the genus Odontaster. All, includ- ing O. validus (Pearse and Bosch, 1986), have pelagic development as far as is known. Consequently, as with echinoids, asteroid clades with nonpelagic development are speciose, but most individuals are not very abundant; those with pelagic development have few species, but indi- viduals of some species can be very abundant. Pearse et al. (1991) and Poulin et al. (2002) suggest that this difference in abundance patterns is due to ecological factors: species with pelagic development colonize and thrive in shallow areas disturbed by ice, while those with nonpelagic de- velopment occur in more stable, deeper habitats, where interspecifi c competition is more intense. Comparing two shallow-water habitats, Palma et al. (2007) found that an ice-disturbed habitat is dominated by O. validus and S. neumayeri, species with planktotrophic development, while a less disturbed habitat is dominated by brooding Abatus agassizii. Brooding is widespread among holothurians in the Southern Ocean. Seventeen of the 41 brooding species of holothurians listed worldwide by Smiley et al. (1991) are found in Antarctic and subantarctic waters. Moreover, 15 of those species are in the order Dendrochirotida, with six each in the genera Cucumaria and Psolus, and brooding by an addition species of Psolus was described by Gutt (1991). In addition, 12 of the brooding species of holothuroids are found in the Scotia Arc area (Pearse and Bosch, 1994). These patterns mirror those seen in echinoids and asteroids. Brooding is also widespread among ophiuroids in the Southern Ocean; Mortensen (1936) estimated that about 50% of the species in Antarctic and subantarctic waters are brooders. Pearse and Bosch (1994) list 33 species of brooding ophiuroids, 21 of which are found in the Sco- tia Arc area. Moreover, most of these species are in the most diverse families in these waters, amphiurids, ophia- canthids, and ophiurids (Hendler, 1991). In contrast to the relatively few speciose genera with brooders in the Southern Ocean, brooding species at lower latitudes are scattered among different genera; Hendler (1991:477) suggests from this difference that there ?may be selection for brooding within clades, rather than a propensity for certain clades to evolve brooding? in the Antarctic ophiu- roid fauna. That is, once brooding is established in a clade, speciation is likely to occur. There has been no phylogenetic analysis of the 12 spe- cies of Southern Ocean crinoids reported to brood, all of them occurring in the Scotia Arc region (Pearse and Bosch, 1994). However, phylogenetic analyses of Promachocrinus kerguelensis in the Atlantic section of the Southern Ocean revealed at least fi ve ?species-level? clades (Wilson et al., 2007). P. kerguelensis is found throughout Antarctic and subantarctic waters, and the one population studied, in McMurdo Sound, produces large numbers of pelagic, leci- thotrophic larvae (McClintock and Pearse, 1987). Find- 13_Pearse_pg181-196_Poles.indd 19013_Pearse_pg181-196_Poles.indd 190 11/17/08 8:37:12 AM11/17/08 8:37:12 AM BROODING AND SPECIES DIVERSITY IN THE SOUTHERN OCEAN 191 ing such cryptic speciation suggests that other populations might brood or have other means of reducing dispersal. CRUSTACEANS Peracarid crustaceans, especially amphipods and iso- pods, are among the most diverse taxa in the Southern Ocean (Held, 2003; Raupach et al., 2004, 2007; L?rz et al., 2007) and are the major contributor to the high spe- cies diversity in those waters. Indeed, the extraordinary species richness of peracarids documented by recent Ant- arctic deep-sea benthic biodiversity (ANDEEP) cruises (Brandt et al., 2004, 2007a, 2007b) in the Atlantic sector of the deep Southern Ocean challenges the idea that a lati- tudinal gradient exists in the Southern Hemisphere. More- over, molecular analyses have revealed additional cryptic species in isopods (Held, 2003; Held and W?gele, 2005; Raupach et al., 2007). All peracarids brood embryos, and most release juveniles that remain close to their parents after being released (the exceptions include exoparasitic isopods, e.g., Dajidae and Bopyridae, and pelagic forms such as mysids and hyperiid amphipods). In contrast to the peracarids, species of decapod crus- taceans, almost all of which release pelagic larvae after brooding embryos, are remarkably few in today?s Southern Ocean. Only a few species of caridean shrimps inhabit the Southern Ocean, and all produce pelagic larvae, even those in the deep sea (Thatje et al., 2005a). Brachyuran crabs are important components of Patagonian benthic ecosystems (Arntz et al., 1999; Gorny, 1999), yet they are entirely ab- sent from the Scotia Arc and Antarctic waters. Recent re- cords of lithodid anomuran crabs in the Southern Ocean indicate a return of these crabs to the Antarctic, perhaps as a consequence of global warming, after their extinction in the lower Miocene ((15 Ma) (Thatje et al., 2005b). The dichotomy in the Southern Ocean between a scar- city of decapods, which have pelagic larvae, and a richness of peracarids, which do not have pelagic larvae, fi ts the ex- tinction hypothesis (Thatje et al., 2005b). Peracarids con- stitute an important part of the prey of lithodid crabs (Co- moglio and Amin, 1999). After the climate deteriorated in the Eocene/Oligocene and benthic decapods became extinct, the absence or scarcity of these top predators may well have created new adaptive zones, leading to a selec- tive advantage for peracarids and favoring their diversi- fi cation. Indeed, free ecological niches may have opened opportunities for spectacular adaptive radiations, as seen in some peracarid taxa (Brandt, 1999, 2005; Held, 2000; L?rz and Brandt, 2004; L?rz and Held, 2004), which were also favored because of their brooding biology. The exceptionally high species diversity of peracarids, especially isopods in the Southern Ocean and its deep en- vironment, cannot, however, be due simply to the fact that they are brooders without pelagic larvae. Peracarids are found throughout the world?s oceans, including the Arctic. However, the Southern Ocean deep-sea samples revealed a strikingly high biodiversity (Brandt et al., 2007a, 2007b). Rather, the high diversity of peracarids in the Southern Ocean may better be accounted for by the unusual ocean- ographic and topographic conditions there, namely, the ACC that has been sweeping through Drake Passage for 30 million years or more. If brooding individuals have been continually displaced by that current and survive downstream in isolation from the parent population, a major ?species diversity pump? would result, producing many species over time. The distribution of species in the well-studied isopod genus Antarcturus reveals the pattern predicted by the ACC hypothesis (Figure 2); 7 of the 15 spe- cies are found in the Scotia Arc? Weddell Sea sector, and an additional 6 are found on the coast of eastern Antarctica. Considering the extensive amount of work that has been done in the Ross Sea during the twentieth century, the bias in species richness toward the Scotia Arc? Weddell Sea and eastern Antarctic coast is unlikely to be a sampling arti- fact. Several other conditions may have contributed to the high diversity of peracarids in the Southern Ocean. Gaston (2000), for example, correlated high habitat heterogeneity with high diversity, and high levels of tectonic activity in FIGURE 2. Distribution of 15 species of Antarcturus around Antarc- tica. Data were compiled from Brandt (1991), Brandt et al. (2007c), and unpublished records from Polarstern expeditions. 13_Pearse_pg181-196_Poles.indd 19113_Pearse_pg181-196_Poles.indd 191 11/17/08 8:37:13 AM11/17/08 8:37:13 AM 192 SMITHSONIAN AT THE POLES / PEARSE ET AL. the Scotia Arc region could have produced relatively high habitat heterogeneity, although not as high as coral reef areas in lower latitudes (Crame, 2000). Mitochondrial gene sequence analyses of iphimediid amphipods, endemic to the Antarctic, indicate that the age of the last common ancestor of this group is approximately 34 million years (L?rz and Held, 2004), after the Southern Ocean was isolated from other fragments of Gondwana- land but well before the Pliocene-Pleistocene glacial sheets extended over the Antarctic Continental Shelf. Speciation, therefore, has probably taken place throughout the time since the ACC became established with the breakthrough of Drake Passage. In summary, crustaceans appear to have patterns of di- versity similar to those seen in echinoderms: relatively few major taxa, which are likely monophyletic clades. Some of the peracarid clades are extremely diverse and speciose, while the decapod clades present, which have pelagic devel- opment, are relatively depauperate in terms of species rich- ness. This pattern indicates that brooding is not so much an adaptation to conditions in the Antarctic but that excep- tional conditions in Antarctic waters enhance speciation of brooders. CONCLUSIONS 1. While nonpelagic development is certainly an ad- aptation resulting from natural selection, it may not be an adaptation to any condition in the present-day Southern Ocean. There is no evidence that nonpelagic development is adaptive to polar conditions or, in particular, to condi- tions in the Southern Ocean. Instead, it may have devel- oped in other environments long ago and is now phyloge- netically constrained. 2. It is possible that most species with lecithotrophic development (pelagic as well as nonpelagic) survived peri- ods when the Antarctic Continental Shelf was largely cov- ered with glacial ice and the Southern Ocean was largely covered with multiyear sea ice, while most species with planktotrophic larvae went extinct because of severely reduced primary production of food for the larvae. The net effect would be (1) an increase in the proportion of species with lecithotrophic development (both pelagic and nonpelagic) and (2) an overall decrease in species richness/ biodiversity. However, the Southern Ocean is notable for its high species richness/diversity. 3. Speciation could be enhanced in taxa with nonpe- lagic development when the following occur: (1) Refuges form on the Antarctic Continental Shelf during the glacial maxima, fragmenting populations into small isolated units that could undergo speciation. If these formed repeat- edly during the glacial-interglacial cycles of the Pliocene- Pleistocene, a ?species diversity pump? would be created. This idea, termed ACS hypothesis, predicts the presence of many closely related cryptic species around the Antarctic continent, mainly at shelf and slope depths. (2) Individuals of species with nonpelagic development are infrequently carried to new habitats by the ACC fl owing through the Drake Passage and over the Scotia Arc, where, if estab- lished, they form new species. Over more than 30 million years, such a process could generate many species. This idea, termed the ACC hypothesis, predicts the existence of many species in clades of varied divergence times, at a wide range of depths but with highest diversity downstream of Drake Passage, in the Scotia Arc and Weddell Sea. 4. All these possibilities appear to be important, de- pending on the taxon of concern, for explaining the un- usual abundance of species with nonpelagic development in the Southern Ocean, but emerging data are giving most support for the ACC hypothesis. In addition, the ACC hy- pothesis may help account for the relatively high diversity found for many taxa in the Southern Ocean, especially in the area of the Scotia Arc and Weddell Sea. ACKNOWLEDGMENTS We are indebted to the superb library of Stanford Uni- versity?s Hopkins Marine Station for making accessible most of the literature reviewed in this paper. Katrin Linse, British Antarctic Survey, Cambridge, provided useful sug- gestions when the manuscript was being developed, and Vicki Pearse, University of California, Santa Cruz, added substantially both to the thinking and content that went into the work as well as with her editorial skills. We thank Andy Clarke, British Antarctic Survey, Cambridge; Ingo Fetzer, UFZ-Center for Environmental Research, Leipzig; Andy Mahon, Auburn University; Chris Mah, Smithson- ian Institution; Jim McClintock, University of Alabama at Birmingham, and anonymous reviewers for comments and information that greatly improved the manuscript. We also thank Rafael Lemaitre, Smithsonian Institution, for inviting us to participate in the Smithsonian International Polar Year Symposium, where the ideas for this synthesis came together. Support for this work was provided by the National Science Foundation (grant OPP-0124131 to JSP) and the German Science Foundation. This is ANDEEP publication number 112. 13_Pearse_pg181-196_Poles.indd 19213_Pearse_pg181-196_Poles.indd 192 11/17/08 8:37:16 AM11/17/08 8:37:16 AM BROODING AND SPECIES DIVERSITY IN THE SOUTHERN OCEAN 193 LITERATURE CITED Absher, T. M., G. Boehs, A. R. Feijo, and A. C. Da Cruz. 2003. 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Villinski, M. Byrne, and R. A. Raff. 2002. Conver- gent Maternal Provisioning and Life-History Evolution in Echino- derms. Evolution, 56: 1764? 1775. Wilson, N. G., R. L. Hunter, S. J. Lockhart, and K. M. Halanych. 2007. Multiple Lineages and Absence of Panmixia in the ?Circumpolar? Crinoid Promachocrinus kerguelensis from the Atlantic Sector of Antarctica. Marine Biology, 152: 895? 904. Wray, G. A. 1995. ?Evolution of Larvae and Developmental Modes.? In Ecology of Marine Invertebrate Larvae, ed. L. McEdward, pp. 413? 447. London: CRC Press. Young, C. M. 1994. ?A Tale of Two Dogmas: The Early History of Deep-Sea Reproductive Biology.? In Reproduction, Larval Biology, and Recruitment in the Deep-Sea Benthos, ed. K. J. Eckelbarger and C. M. Young, pp. 1? 25. New York: Columbia University Press. 13_Pearse_pg181-196_Poles.indd 19613_Pearse_pg181-196_Poles.indd 196 11/17/08 8:37:19 AM11/17/08 8:37:19 AM ABSTRACT. Trawl surveys conducted between 1996 and 2007 show that populations of octopods have signifi cantly higher abundances around Elephant Island, off the Ant- arctic Peninsula, than in similar areas nearby. This elevated abundance was fi rst detected following the cessation of commercial fi shing and has persisted for many years beyond, possibly indicating an enduring shift in the structure of the ecosystem. INTRODUCTION Concern about the effects of overfi shing on marine ecosystems has increased substantially in recent years (e.g., Jackson et al., 2001). One of these potential effects is a shift in the suite of dominant predators in the ecosystem (Fogarty and Murawski, 1998; Choi et al., 2004). Unusually high abundances of squids and octopods in some areas have been related to man?s removal of their fi nfi sh predators and competitors (Caddy and Rodhouse, 1998). Furthermore, anthro- pogenic changes in polar regions are of particular conservation concern (e.g., Smith et al., 2002). In Antarctica, a bottom-trawl fi shery primarily targeting mackerel icefi sh (Champsocephalus gunnari) and marbled notothenia (Noto- thenia rossii) developed in 1978 around Elephant Island, in the South Shetland Archipelago off the Antarctic Peninsula. The fi shery continued until 1988/1989 but rapidly depleted the populations of the target species (Kock and Stransky, 2000). We report here that in this overfi shed area, populations of octopods have signifi cantly higher abundances than in similar areas nearby. This elevated abun- dance has persisted for years beyond the cessation of commercial fi shing, pos- sibly indicating an enduring shift in the structure of the ecosystem. MATERIALS AND METHODS R/V Polarstern cruises ANT XIV/2 (November? December 1996), ANT XIX/ 3 (January? February 2002), and ANT XXIII/8 (December 2006 to January 2007) were conducted to assess the status of fi sh stocks in the region around Ele- phant Island monitored internationally under the Convention on Conservation Michael Vecchione, National Marine Fisheries Service, Systematics Laboratory, National Mu- seum of Natural History, P.O. Box 37012, MRC 153, Washington, DC 20013-7012, USA. Louise Allcock, Martin Ryan Marine Science Institute, National University of Ireland, Galway, Univer- sity Road, Galway, Ireland. Uwe Piatkowski, Institute for Marine Research, Universitat Kiel, Dusternbrooker Weg 20, D-24105, Kiel, Ger- many. Elaina Jorgensen, National Marine Fisher- ies Service, Alaska Fisheries Science Center, 7600 Sand Point Way N.E., Seattle, WA 98115, USA. Iain Barratt, Ecology and Evolutionary Biology, School of Biological Sciences, Queen?s University Belfast, 97 Lisburn Road, Belfast BT9 7BL, UK. Corresponding author: M. Vecchione (vecchiom@ si.edu). Accepted 19 May 2008. Persistent Elevated Abundance of Octopods in an Overfi shed Antarctic Area Michael Vecchione, Louise Allcock, Uwe Piatkowski, Elaina Jorgensen, and Iain Barratt 14_Vecchione_pg197-204_Poles.ind197 19714_Vecchione_pg197-204_Poles.ind197 197 11/18/08 9:16:01 AM11/18/08 9:16:01 AM 198 SMITHSONIAN AT THE POLES / VECCHIONE ET AL. of Antarctic Marine Living Resources. Sampling stations were selected randomly from depth strata between 50 and 500 m. These stations were sampled by 30-min tows with a large double-warp otter trawl. The 2002 cruise also con- ducted similar sampling off the southern South Shetland Islands and off Joinville Island across the Bransfi eld Strait (both areas with shelves of similar widths and depths to the Elephant Island area), as well as an intensive sampling series of 20 tows in a shallow-water grid near Elephant Is- land. The 2006? 2007 cruise similarly sampled additional stations across the Bransfi eld Strait close to the peninsula and in the western Weddell Sea (Figure 1). We identifi ed and counted all cephalopods collected on these cruises, in- cluding both the cod end sample and specimens entangled in the net mesh. The material included many more spe- cies than were recognized previous to this work (Allcock, 2005). We fi rst noticed the abundance patterns reported here during the 2002 cruise and have since examined the 2006? 2007 cruise as a test of our unpublished 2002 hy- pothesis. Because the sample sizes, depths, etc., were bal- anced between the Elephant Island area and the out-groups in 2002, we emphasize these results and present the 1996 and 2006? 2007 results for temporal comparisons. For some of the comparisons presented below, we have eliminated the shallow-grid samples and pooled the obser- vations from the southern South Shetland Islands, close to the Antarctic Peninsula, Joinville Island, and Weddell Sea (termed ?out-groups? below) because this created similar- sized sets of samples with similar ranges and variances in depth, a controlling factor in the abundance and diversity of Antarctic octopods (Figure 2). Intensive shallow-grid sampling was not conducted in the out-group areas. A FIGURE 1. Study area showing sampling locations. 14_Vecchione_pg197-204_Poles.ind198 19814_Vecchione_pg197-204_Poles.ind198 198 11/18/08 9:16:02 AM11/18/08 9:16:02 AM ABUNDANCE OF ANTARCTIC OCTOPODS 199 FIGURE 2. Relationship between mean depth of tow and octopod catch in 2002 trawl samples from around Elephant Island and similar nearby areas: (a.) total number of octopods per tow, (b.) approximate number of octopod species per tow. 14_Vecchione_pg197-204_Poles.ind199 19914_Vecchione_pg197-204_Poles.ind199 199 11/18/08 9:16:54 AM11/18/08 9:16:54 AM 200 SMITHSONIAN AT THE POLES / VECCHIONE ET AL. summary of the collections is presented in Table 1. Statisti- cal comparisons used two-sample t-tests assuming unequal variances, with the a priori confi dence level for signifi cance at H9251 H11005 0.05 (two tailed). Depths presented are based on the mean depth of each tow, calculated as the average of the depth at the beginning and at the end of the tow. RESULTS Although the depths of the 28 Elephant Island sta- tions sampled in 2002 were slightly shallower than the 26 out-group stations, the difference was not statistically sig- nifi cant. However, the total number of octopods collected per tow at Elephant Island averaged over twice as high (Table 2) as that at the out-group stations (signifi cant, p H11021 0.001). Had we included the 20 shallow-grid stations, the difference in catch between the areas would have been even greater (Table 1) because these tows included two catches that were anomalously high (98 and 306 octopods) for that depth range and strongly dominated by one shallow- dwelling species, Pareledone charcoti (Joubin, 1905). Al- though the difference between areas in number of species collected was not great, it was statistically signifi cant. This is likely because an increased number of specimens in the catch generally includes a higher number of species (Figure 3) rather than because of a difference in species richness in the two areas. Depths of the 38 stations sampled around Elephant Island in 1996 were not signifi cantly different from those sampled in 2002, nor was the average catch signifi cantly different from the 2002 catch in the same area. Only three samples were collected in 1996 from the southern South Shetland Islands (in the same area as the out-groups in 2002) with the same net at similar depths; the catches in these samples were very low (12, 7, and 5 octopods). The few samples that were collected from Joinville Island in 2002 included the lowest numbers of octopods caught that year, but those were qualitatively similar to the octopod fauna of the Weddell Sea (Allcock et al., 2001). In 2006? 2007, both the mean depth and number of octopods per sample around Elephant Island were less than in 2002. The mean depth at the out-group stations in 2006? 2007 was somewhat greater than in 2002 and signifi cantly greater than the Elephant Island stations in 2006? 2007. However, the number of octopods per sample around Ele- phant Island was H110222.5 times higher than in the out-group samples, a statistically signifi cant difference (Table 3). As in TABLE 1. Summary of octopod collections by cruise year and location. Number 95% confi dence Year Location of stations Parameter a Maximum Mean interval Minimum 2006? 2007 Elephant Island 51 Depth (m) 486 208 26 62 catch 169 18 8 0 no. spp. 8 3 0.5 0 2006? 2007 out-group 38 depth (m) 490 275 35 87 catch 35 7 2 0 no. spp. 5 2 0.5 0 2002 Elephant Island 28 depth (m) 455 257 32 127 catch 145 42 16 1 no. spp. 9 6 1 1 2002 shallow grid 20 depth (m) 209 146 18 74 catch 306 37 15 4 no. spp. 7 4 1 2 2002 out-group 26 depth (m) 468 266 49 85 catch 80 18 10 1 no. spp. 10 4 1 1 1996 Elephant Island 38 depth (m) 477 243 33 89 catch 135 52 10 9 a Depth is based on mean depth of each tow, catch is the number of octopods per tow, and no. spp. is the approximate number of species per tow; no. spp. is not included for 1996 cruise because identifi cations have not yet been revised based on updated taxonomy resulting from 2002 cruise. 14_Vecchione_pg197-204_Poles.ind200 20014_Vecchione_pg197-204_Poles.ind200 200 11/18/08 9:16:57 AM11/18/08 9:16:57 AM ABUNDANCE OF ANTARCTIC OCTOPODS 201 FIGURE 3. Relationship between number of octopods collected and approximate number of species collected in 2002. 2002, the larger catches of octopods around Elephant Is- land included a signifi cantly greater number of species than in the smaller out-group samples. DISCUSSION We have no quantitative information about the abun- dance of octopods in the area of Elephant Island prior to the onset of commercial fi shing in the area. Therefore, some unknown natural factor in that location could have resulted in high octopod abundances relative to levels in similar areas nearby. However, at least eight bottom-trawl surveys of the fi sh fauna around Elephant Island were con- ducted between 1976 and 1987 (Kock and Stransky, 2000), and we know of no indication of elevated octopod abun- dance in those surveys. Conversely, during a U.S. trawl survey in 1998, octopod abundance was higher around Elephant Island than off the South Shetland Islands (C. Jones, NMFS Antarctic Program, Southwest Fisheries Sci- ence Center, personal communications, 2002). These sam- pling efforts, together with the present surveys, revealed that bycatch fi sh species had recovered by the early 1990s but populations of target species had not fully recovered (Kock and Stransky, 2000; K.-H. Kock, Institute for Sea Fisheries, personal communication, 2002). The abun- dance of octopods may have increased coincidently with the recovery of populations of fi nfi sh bycatch. We are not sure why the overall abundance of octo- pods around Elephant Island was lower in 2006? 2007 than during previous surveys. Perhaps this is because the sampling was concentrated at somewhat shallower depths, which resulted in fewer catches with very high numbers of octopods (Figure 2). However, details of the confi guration of the trawl, such as height of the footrope above the roll- ers and the presence of a ?tickler? chain, were modifi ed during this cruise to reduce the bycatch of sessile mega- fauna (e.g., sponges and cnidarians). It seems quite likely that these modifi cations affected the net?s sampling char- acteristics for benthic octopods. An indication that high numbers of octopods remained around Elephant Island during 2006? 2007 is found in the very high numbers of small specimens caught at three stations (59 specimens at station 614-3, 24 at 642-1, and 15 at 654-6) using a differ- ent gear type, an Agassiz beam trawl, not included in the comparisons presented here. The pattern reported here is consistent with other re- ports from around the world of elevated cephalopod abun- dances coincident with reduction of fi nfi sh populations by commercial harvesting. As the inferred increase in octo- pod abundance appears to be coincident with recovery of bycatch fi sh populations, release from predation pressure 14_Vecchione_pg197-204_Poles.ind201 20114_Vecchione_pg197-204_Poles.ind201 201 11/18/08 9:16:58 AM11/18/08 9:16:58 AM 202 SMITHSONIAN AT THE POLES / VECCHIONE ET AL. rather than competitive processes seems to be responsible for the good fortune of the octopods. Because ecological processes in polar regions tend to be comparatively slow, such ecosystem-level impacts of fi shing may take longer to become apparent than at lower latitudes but also may be very persistent. ACKNOWLEDGMENTS We thank Karl-Hermann Kock and the Institute for Sea Fisheries in Hamburg, Germany, for allowing us to par- ticipate in these cruises, which were conducted by the Al- fred Wegener Institute for Polar Research in Bremerhaven, Germany. Silke Stiemer (Universitat Kiel) assisted with the fi eld work, and R. E. Young (University of Hawaii) and C. F. E. Roper (Smithsonian Institution) provided helpful comments on an early draft of the manuscript. LITERATURE CITED Allcock, A. L. 2005. On the Confusion Surrounding Pareledone charcoti: Endemic Radiation in the Southern Ocean. Zoological Journal of the Linnean Society, 143(1): 75? 108. Allcock, A. L., U. Piatkowski, P. K. G. Rodhouse, and J. P. Thorpe. 2001. A Study on Octopodids from the Eastern Weddell Sea, Antarctica. Polar Biology, 24: 832? 838. Caddy, J. F., and P. G. Rodhouse. 1998. Cephalopod and Groundfi sh Landings: Evidence for Ecological Change in Global Fisheries? Re- views in Fish Biology and Fisheries, 8: 431? 444. Choi, J. S., K. T. Frank, W. C. Leggett, and K. Drinkwater. 2004. Transi- tion to an Alternate State in a Continental Shelf Ecosystem. Cana- dian Journal of Fisheries and Aquatic Sciences, 61:505? 510. TABLE 2. Two-sample t-test assuming unequal variances comparing areas in February 2002. a Parameter Elephant Island Out-groups Station depths Mean 256.9892857 265.8461538 Variance 6694.207844 14457.90058 Observations 28 26 df 44 t statistic H110020.314090727 P(T H11349 t) one-tail 0.377468272 t critical one-tail 1.680230071 P(T H11349 t) two-tail 0.754936545 t critical two-tail 2.0153675 Number of octopods per tow Mean 42.85714286 18.07692308 Variance 1652.941799 609.6738462 Observations 28 26 df 45 t statistic 2.728499663 P(T H11349 t) one-tail 0.004521735 t critical one-tail 1.679427442 P(T H11349 t) two-tail 0.009043469 t critical two-tail 2.014103302 Number of species per tow Mean 5.535714286 4 Variance 6.406084656 7.36 Observations 28 26 df 51 t statistic 2.146507581 P(T H11349 t) one-tail 0.018305786 t critical one-tail 1.675284693 P(T H11349 t) two-tail 0.036611573 t critical two-tail 2.007582225 a Out-groups are pooled samples from southern South Shetland Islands plus Joinville Island. On the basis of these analyses, the depths of the stations sampled do not differ signifi cantly between the areas, but both the number of octopods per tow and the number of octopod species per tow were signifi cantly higher in the area around Elephant Island than in other areas. 14_Vecchione_pg197-204_Poles.ind202 20214_Vecchione_pg197-204_Poles.ind202 202 11/18/08 9:17:01 AM11/18/08 9:17:01 AM ABUNDANCE OF ANTARCTIC OCTOPODS 203 TABLE 3. Two-sample t-test assuming unequal variances comparing areas in December 2006 to January 2007. a Parameter Elephant Island Out-groups Station depths Mean 208.2411765 274.6736842 Variance 8640.174871 11619.0328 Observations 51 38 df 73 t statistic H110023.047557645 P(T H11349 t) one-tail 0.001605534 t critical one-tail 1.665996224 P(T H11349 t) two-tail 0.003211069 t critical two-tail 1.992997097 Number of octopods per tow Mean 18.23529412 7.184210526 Variance 733.5835294 45.66785206 Observations 51 38 df 58 t statistic 2.799243278 P(T H11349 t) one-tail 0.003472082 t critical one-tail 1.671552763 P(T H11349 t) two-tail 0.006944163 t critical two-tail 2.001717468 Number of species per tow Mean 3.215686275 2.473684211 Variance 3.77254902 1.877667141 Observations 51 38 Hypothesized mean difference 0 df 87 t statistic 2.11239907 P(T H11349 t) one-tail 0.018759109 t critical one-tail 1.66255735 P(T H11349 t) two-tail 0.037518218 a Out-groups are pooled samples from areas other than Elephant Island. In these analyses, the number of octopods per tow and the number of octopod species per tow were signifi cantly higher in the area around Elephant Island than in other areas, but the sampled depths around Elephant Island were shallower, which could be a confounding factor. Fogarty, M. J., and S. A. Murawski. 1998. Large-Scale Disturbance and the Structure of Marine Systems: Fishery Impacts on Georges Bank. Ecological Applications, 8(S1): S6? S22. Jackson, J., M. X. Kirby, W. H. Berger, K. A. Bjorndal, L. W. Botsford, B. J. Bourque, R. H. Bradbury, R. Cooke, J. Erlandson, J. A. Estes, T. P. Hughes, S. Kidwell, C. B. Lange, H. S. Lenihan, J. M. Pandolfi , C. H. Peterson, R. S. Steneck, M. J. Tegner, and R. R. Warner. 2001. Historical Overfi shing and the Recent Collapse of Coastal Ecosys- tems. Science, 293: 629? 638. Kock, K.-H., and C. Stransky. 2000. The Composition of the Coastal Fish Fauna around Elephant Island (South Shetland Islands, Ant- arctica). Polar Biology, 23: 825? 832. Smith, J., R. Stone, and J. Fahrenkamp-Uppenbrink. 2002. Trouble in Polar Paradise. Science 297: 1489. 14_Vecchione_pg197-204_Poles.ind203 20314_Vecchione_pg197-204_Poles.ind203 203 11/18/08 9:17:02 AM11/18/08 9:17:02 AM 14_Vecchione_pg197-204_Poles.ind204 20414_Vecchione_pg197-204_Poles.ind204 204 11/18/08 9:17:02 AM11/18/08 9:17:02 AM ABSTRACT. Antarctic bryozoans are spectacular. They are often larger and more col- orful than their temperate relatives. Antarctic bryozoans are also outstanding in their diversity. Well over 300 species have been described, and new descriptions continue to appear. In the U.S. Antarctic Research Program (USARP) collections we have identifi ed 389 species so far, mostly belonging to the Cheilostomata, the dominant order in Recent seas. Much about their ecology can be learned from study of the abundant material preserved in the USARP collections. Yearly growth bands demonstrate that colonies may live for decades and that growth rates are very close to those of related temperate species. The presence of embryos in the brood chambers of many species allows determination of seasonality of reproduction and fecundity of colonies of different sizes. A large propor- tion of Antarctic bryozoan species (81% for cheilostomes) are endemic. Endemic groups include bizarre and unusual forms in which polymorphism, the occurrence of individuals specialized to perform different tasks, is highly developed. Behavioral studies carried out with living colonies in the Antarctic have shown how different polymorphs function in cleaning and protecting colonies from trespassers or predators: capturing motile animals such as amphipods, polychaetes, and nematodes and sweeping colonies free of debris. INTRODUCTION The more than 5,000 members of the phylum Bryozoa, belonging to the Lophotrochozoa group of protostome invertebrates, are found in marine and freshwater habitats worldwide, including the seas that surround Antarctica. Bryozoans are colonial, benthic, sessile animals. They reproduce asexually by budding new members of a colony or, in some cases, by fragmentation of an ex- isting colony. Colonies also reproduce sexually. Embryos, often brooded, develop into free-swimming larvae that settle and metamorphose to begin a new colony. Although individual bryozoan zooids are microscopic (ranging from about 0.30 to 2.0 mm in length), colonies can be quite large, consisting of many thousands of individuals and, in some places, creating three-dimensional benthic habitats that serve as shelter, feeding, and nursery grounds for other organisms. Bryozo- ans are suspension feeders, part of the benthic biological fi lter system made up of sessile fi lter-feeding animals. Bryozoans also produce physical and chemical defenses against their enemies. Many members of the dominant Recent order of bryozoans, the cheilostomes, have developed a high degree of polymorphism, Judith E. Winston, Department of Marine Biol- ogy, Virginia Museum of Natural History, 21 Starling Avenue, Martinsville, VA 24112, USA (judith.winston@vmnh.virginia.gov). Accepted 19 May 2008. Cold Comfort: Systematics and Biology of Antarctic Bryozoans Judith E. Winston 15_Winston_pg205-222_Poles.indd 20515_Winston_pg205-222_Poles.indd 205 11/17/08 9:21:54 AM11/17/08 9:21:54 AM 206 SMITHSONIAN AT THE POLES / WINSTON with heterozooids specialized for reproduction, attach- ment, cleaning, and defense. Their chemical defenses con- sist of natural products such as alkaloids or terpenoids, which are toxic or discouraging to predators or disease agents (Sharp et al., 2007). The chilly Antarctic seafl oor, where seawater tempera- tures are near or below freezing year-round, might not seem like an ideal habitat for benthic organisms. Food is plentiful for only part of the year, and, in shallow water, icebergs may scour the seafl oor, destroying everything they touch. Yet more than 300 species of bryozoans have been found there, including a large number of distinctive endemic taxa. This paper will discuss some results from more than 20 years of study of the U.S. Antarctic Research Program (USARP) bryozoan collections by the author and collaborators A. E. Bernheimer, P. J. Hayward, and B. F. Heimberg. TAXONOMY AND DISTRIBUTION OF USARP BRYOZOANS BRYOZOAN COLLECTION IN THE ANTARCTIC The beginning of bryozoan collecting in the Antarctic dates from the last part of the nineteenth century and the early part of the twentieth, the era of the national expedi- tions for the exploration of the continent. Although a few species from Cape Adare in the Ross Sea were recorded by Kirkpatrick (1902), the fi rst major report on the bryozoan fauna was by Waters (1904) on the 89 taxa collected in the Bellingshausen Sea by the Belgica Expedition. Analy- sis of collections from other expeditions followed (Calvet, 1904a, 1904b, 1909; Kluge, 1914; Thornely, 1924; Liv- ingstone, 1928). A second wave of collecting began with a series of voyages carried out by the British Discovery In- vestigations (of which some of the bryozoans appeared in Hastings, 1943). The fi rst U.S. specialist to study Antarctic bryozoans was Mary Rogick, who published 13 papers on the taxonomy of the bryozoans collected during the U.S. Navy?s 1947? 1948 Antarctic Expedition between 1955 and 1965 (summarized in Rogick, 1965). THE USARP BRYOZOAN COLLECTIONS The International Geophysical Year of 1957? 1958 marked a new era of marine research and benthic col- lecting in the Antarctic. The USARP bryozoan collections consist of more than 5,000 lots collected by various U.S. research groups between 1958 and 1982. The earliest lots in the collection were obtained during benthic studies by Stanford University researchers working in the McMurdo Sound and the Ross Sea. Later systematic collections of all benthos, including bryozoans, were made by scientists and technicians during oceanographic cruises of USNS Eltanin (1962? 1968) and R/V Hero (1968? 1982). These collections were returned to the United States to be pro- cessed by technicians at the Smithsonian Oceanographic Sorting Center (SOSC) and were distributed to taxonomic specialists on each group. The stations from which bryo- zoans were collected ranged from 10?W to 70?E but were concentrated in the following Antarctic and subantarctic areas: the Ross Sea, the Antarctic Peninsula, off the islands of the Scotia Arc, and from Tierra del Fuego to the Falk- land Islands (Figure 1). My involvement with the collections included training SOSC technicians in bryozoan identifi cation and working with technicians and volunteers at SOSC, the American Museum of Natural History, and the Virginia Museum of Natural History to sort and identify those lots, now sepa- rated into almost 8,000 vials and jars. It also led to a 1985 NSF grant to work at Palmer Station to study ecology and behavior of living colonies of some Antarctic species. In spite of the large amount of marine biology carried out in Antarctica, relatively little attention had been paid to benthic community structure or to the ecology of benthic organisms. For this reason, as part of the project on be- havioral and chemical ecology of Antarctic bryozoans, FIGURE 1. Location of USARP stations containing bryozoans pro- cessed by the Smithsonian Oceanographic Sorting Center. 15_Winston_pg205-222_Poles.indd 20615_Winston_pg205-222_Poles.indd 206 11/17/08 9:21:54 AM11/17/08 9:21:54 AM SYSTEMATICS AND BIOLOGY OF ANTARCTIC BRYOZOANS 207 we attempted to learn more about the role these animals played in one particular benthic community, that off the southern end of Low Island, South Shetland Islands. Finally, as a lecturer on the American Museum?s Discov- ery tours, I made three additional trips to the Antarctic Peninsula and South Shetland Islands, where I was able to observe beach drift organisms and intertidal bryozoan habitats at a number of locations. TAXONOMIC DISTRIBUTION OF USARP BRYOZOANS Two classes, Gymnolaemata and Stenolaemata, and three orders of bryozoans are recorded from the Antarc- tic. The Gymnolaemata include the orders Ctenostomata and Cheilostomata. The Cyclostomata are the only living order of the Stenolaemata. The dominant bryozoans in Recent oceans are the cheilostomes, a group whose diversity and abundance has increased greatly since its origin in the Cretaceous. Chei- lostomes have box-like zooids, variously reinforced with calcium carbonate skeletons, a hinged operculum pro- tecting the orifi ce through which the polypide extends to feed, and colonies often displaying a high degree of poly- morphism. Most cheilostomes brood their embryos to the larval stage in variously formed reproductive structures called ovicells. Of the 389 species so far recorded in the sorted and identifi ed USARP collections, 344 (88.5%) are cheilo- stomes. Four new species have been described so far from the collection (Hayward and Winston, 1994) with more to follow. Although most of the major groups of cheilo- stomes are represented in the Antarctic and subantarctic, several groups have undergone a good deal of radiation with numerous species, most of them endemic. Families especially well represented (10 or more species) include the Flustridae, Calloporidae, Bugulidae, Cabereidae, Cel- lariidae, Arachnopusiidae, Exochellidae, Sclerodomidae, Smittinidae, Microporellidae, Celleporidae, and Retepo- ridae (Hayward, 1995). This pattern is refl ected in the USARP collections. Some representative species are shown in Figure 2. Although cyclostome bryozoans rank well below chei- lostomes in terms of species diversity, they are abundant in the Antarctic, forming large colonies, easily recogniz- able as cyclostomes, although often diffi cult or impos- sible to determine to species level because colonies lack gonozooids, a vital taxonomic character in the group. Like cheilostomes, cyclostome bryozoans have calcifi ed walls. Their zooids are tubular. The zooid orifi ce, at the distal end of the tube, has no operculum but closes by a sphinc- ter instead. Polymorphism is much less common, but some species have nanozooids. They brood embryos in one or more brood chambers or gonozooids, and sexual repro- duction is unusual, involving polyembryony: larval fi ssion in which the original fertilized egg results in many geneti- cally identical larvae. Thirty-seven cyclostomes (9.55% of the total USARP bryozoan fauna) were found in USARP collections. Figure 3 (left) shows an erect branching cyclo- stome of the genus Hornera. Members of the order Ctenostomata have chitinous and/or gelatinous walls. Some form encrusting or massive colonies. Others form delicate vine-like colonies consisting of single or clumps of tubular zooids along stolons or with zooid bases constricting into stolonate tubes that join adja- cent zooids. They have no operculum but close the orifi ce by muscular constriction. Sexual reproduction may involve brooding embryos in body cavities of maternal zooids or broadcasting fertilized eggs into seawater, where develop- ment into larvae takes place. Although the fi rst bryozoan described from the Antarctic continent was a ctenostome, Alcyonidium fl abelliforme (Kirkpatrick, 1902), in terms of species diversity the group is less well represented there than in some other environments. Only eight species (2%) of the total bryozoan fauna were found in the USARP collections, but three of them represented new species (Winston and Hayward, 1994), one encrusting on other bryozoans and the other two boring into live bryozoan colonies or living in dead zooids or crevices (e.g., Bowerbankia antarctica, Figure 3). The most abundant species in the Antarctic Peninsula and South Shetland Islands form foliaceous, lightly calci- fi ed colonies (Carbasea ovoidea, Kymella polaris, Flustra astrovae, Himantozoum antarcticum, and Nematofl ustra fl agellata), delicate jointed branching colonies (Cellaria divisa), or encrusting colonies (e.g., Micropora brevis- sima, Inversiula nutrix, Celleporella antarctica, Ellisina antarctica, Harpecia spinosissima, Lacerna hosteensis, Escharoides tridens, and Lichenopora canaliculata). From the Ross Sea, the most abundant species in the USARP collections in shallow water (less than 50 m) is Eminoecia carsonae, with rigidly branching erect colonies. In deeper water, erect forms are also common, including Cellaria monilorata, Arachnopusia latiavicularis, Antarcticaetos bubeccata, Thryptocirrus phylactellooides, Melicerita obliqua, species of Cellarinellidae, Reteporella antarctica and other Reteporidae, with well-calcifi ed branching or reticulate colony forms (see Figure 4), as well as some of the same epizoic species found in the Antarctic Peninsula area: Celleporella antarctica, Ellisina antarctica, and Har- pecia spinosissima. 15_Winston_pg205-222_Poles.indd 20715_Winston_pg205-222_Poles.indd 207 11/17/08 9:21:55 AM11/17/08 9:21:55 AM 208 SMITHSONIAN AT THE POLES / WINSTON The Antarctic intertidal zone has been considered to be almost barren due to scouring by seasonal sea ice (Barnes, 1994a; Knox, 2007). This is not the case, how- ever, in many localities along the Antarctic Peninsula and in the South Shetlands. Sheltered intertidal sites, often with plankton-rich eutrophic water due to runoff from adjacent colonies of birds or marine mammals, contain tide pools and crevices whose inhabitants include calcar- eous and macroalgae, limpets, amphipods, Glyptonotus, brittle stars, starfi sh, and sea urchins. A number of bryo- zoans, of which Inversiula nutrix (Figure 2d) was most common, encrust intertidal rocks or attach to seaweed in intertidal to shallow subtidal habitats (Winston and Hayward, 1994). FIGURE 2. Morphologies common in bryozoans from USARP collections. (a) Large foliaceous colony of Carbasea ovoidea from Low Island. Scale H11005 5 cm. Arrow points to pycnogonid. (b) Rigid branching colony of Cellarinella foveolata attached to glacial pebble. (c) Living colony of Kymella polaris pinned out for study of reproduction and fouling. (d) Encrusting colonies of Inversiula nutrix on underside of an intertidal rock, shown by arrows. Branching hydroid above colonies and crustose algae (dark patches) are also visible. 15_Winston_pg205-222_Poles.indd 20815_Winston_pg205-222_Poles.indd 208 11/17/08 9:21:55 AM11/17/08 9:21:55 AM SYSTEMATICS AND BIOLOGY OF ANTARCTIC BRYOZOANS 209 BIOLOGY AND ECOLOGY OF ANTARCTIC BRYOZOANS STUDY AREAS, METHODS, AND MATERIALS In addition to long-term taxonomic projects, other studies using preserved USARP material, ecological and behavioral work on living Antarctic bryozoans, were car- ried out during the austral summer and fall of 1985 at Palmer Station and off the southernmost of the South Shetland Islands. Low Island (latitude 63?25H11032S, longitude 62?10H11032W) lies off the western coast of the Antarctic Pen- insula. The Low Island area was a favorite collecting site for fi sh biologists working out of Palmer Station during our stay. It was close enough to the station (within 12 hours? cruise time) that fi sh could be maintained in good condition, and the sea bottom off its southern side slopes off to a relatively fl at surface, free of projecting ledges or pinnacles that would tear the trawls used to collect speci- mens. By sharing cruise time with fi sh and krill biologists we were able to return to the site several times during the summer and fall. A 4 H11003 10 ft (1.2 H11003 3 m) otter trawl was used to collect both fi sh and bryozoans. Proportionate bio- mass of benthic organisms was determined on shipboard by averaging the blotted wet weights of different groups of organisms taken in three timed trawls of equal length. For studies of the food of the bryozoans we froze a number of freshly trawled colonies of the most common species in the ultracold freezer on the ship. They were later defrosted in the lab, and polypides were dissected and gut contents were examined by light and epifl uorescence microscopy. For other studies of living material the bryozoans collected were maintained in holding tanks with running seawater until the ship returned to Palmer Station. On return to the lab they were placed in running seawater tanks and examined as soon as possible. Specimens for behavioral studies were collected both from Low Island by trawling and from Arthur Harbor, using a hand dredge from an infl atable boat. Autozooid feeding and behavior of avicu- larian polymorphs was recorded using a macrovideo setup in the lab at Palmer Station. Freshly collected material was maintained at close to normal seawater temperature dur- ing behavioral observations by an ice bath surrounding the observation dish. To analyze growth and injury, colonies were pinned out in seawater in a shallow dissecting tray. A piece of clear Mylar fi lm was then placed over them, and the col- ony outline, including growth checks, was traced with a waterproof marking pen. The traced version was immedi- ately copied and used as a map to record areas of injury and fouling. Each colony was examined under the dis- secting microscope, and all injuries, discoloration, bites, rips, empty zooids, and fouling organisms were recorded. FIGURE 3. (left) An Antarctic cyclostome, Hornera sp. (scanning electron microscope image). (right) Living colony of an Antarctic ctenostome, Bowerbankia antarctica. Arrow points to lophophore emerging from crevice between two spirorbid tubes. Tentacle crown of spirorbid is visible below that of bryozoan. 15_Winston_pg205-222_Poles.indd 20915_Winston_pg205-222_Poles.indd 209 11/17/08 9:22:01 AM11/17/08 9:22:01 AM 210 SMITHSONIAN AT THE POLES / WINSTON Growth of individual colonies was analyzed by measuring the distance between yearly growth checks. For the reproductive study fi ve colonies (if possible) were examined after each collection. Each colony was pinned out fl at in a dissecting tray fi lled with seawater. It was then examined under a dissecting microscope for the following: developing ovicells, developing embryos, ma- ture embryos, and empty ovicells. The colonies were also photographed and fi nally preserved in 70% ethanol. BIOLOGICAL CHARACTERISTICS Outstanding characteristics of invertebrates occurring in Antarctic benthic communities include a high degree of endemism, large body size in comparison with temper- ate or tropical relatives, and the prevalence of suspension feeding organisms (Hedgpeth, 1969, 1970; Dell, 1972; Arnaud, 1974; White, 1984; Gallardo, 1987; Arntz et al., 1994, 1997; Clarke and Johnston, 2003; Knox, 2007). Many Antarctic bryozoans are large in size. Figure 2a shows a large colony of Carbasea ovoidea, a fl ex- ible, lightly calcifi ed, foliaceous species collected at Low Island, South Shetland Islands. The size of the colony is approximately 20 cm in width by 15 cm in height. Indi- vidual zooids of many Antarctic species also are large in size, many of them between 1 and 2 mm long, versus the more common 0.4? 0.9 mm zooid length of species found in warmer waters. Figures 2 and 3 show some of the range of colony morphology found in species in the USARP collections. Branching colonies consisting of rooted seaweed-like fronds, wide or narrow, are abundant, especially in the Antarctic Peninsula and South Shetland Islands. Jointed and rigid erect forms, branching, unbranched, or anasto- mosing and reticulate, may be attached to other substrata, such as glacial pebbles (Figure 2b), vertical walls, or the dead colonies of other bryozoans or rooted in sediment (Figure 2c). Encrusting species are also abundant and di- verse. Some form massive or nodular colonies consisting of several layers of frontally budded zooids. Other spe- cies form single-layered crusts, loosely or tightly attached to other bryozoans, other organisms, or hard substrata ( Figure 2d). Many Antarctic bryozoans are also more colorful than their temperate relatives. Bryozoans derive their pigment from the carotenoids in their phytoplankton food or, in some cases, from coloration present in symbiotic bacteria inhabiting zooids (Sharp et al., 2007). Colors of living col- onies range from dark red (e.g., Carbasea curva) to orange brown (e.g., Nematofl ustra fl agellata), to yellow orange (e.g., Kymella polaris), to peach (e.g., Orthoporidra spp.), purple, pink (e.g., chaperiids and reteporids), and tan to white. The one Bugula species known, Bugula longissima, has dark green coloration when living, apparently derived from bacterial symbionts (Lebar et al., 2007). The dark red Carbasea curva, which lacks the physical defense of avicularia, was found to show moderate hae- molytic activity, killing 60% of human and 50% of dog erythrocytes (Winston and Bernheimer, 1986). Some of the species from Low Island were also tested for antibiotic ac- tivity. Kymella polaris and Himantozoum antarcticum both strongly inhibited the growth of Staphylococcus aureus. Nematofl ustra fl agellata, Caberea darwinii, and Austro- fl ustra vulgaris moderately inhibited growth. Only Beania livingstonei was noninhibitory (Colon-Urban et al., 1985). STUDIES OF BRYOZOANS FROM THE LOW ISLAND BENTHIC COMMUNITY Shallow shelf environments (less than 500 m) in many areas of the Antarctic are dominated by communities made up largely of sessile suspension feeders like sponges, bryozoans, hydroids, gorgonians, and tunicates, whose colonies may form dense thicket-like growths spreading over large areas of the sea bottom (Belyaev, 1958; Uscha- kov, 1963; Propp, 1970; White and Robins, 1972; Barnes, 1995b, 1995c; Sa?z-Salinas et al., 1997; Sa?z-Salinas and Ramos, 1999; San Vincente et al., 2007). In contrast to epifaunal communities elsewhere, which are mostly lim- ited to hard substrata, such communities in Antarctica commonly rest on or are rooted in soft sediments or are attached to scattered rocks and pebbles. Gallardo (1987) fi rst called attention to the need to for recognizing the dis- tinctiveness of this epi-infaunal or soft-bottom epifaunal community as a prerequisite to the study of its structure. The community we studied fi t this epi-infaunal pattern. Biomass Inshore, in depths of 70 m or less, benthic biomass con- sisted primarily of macroalgae and echinoderms, a com- munity similar to that reported at a number of localities along the peninsula (Gruzov and Pushkin, 1970; Delaca and Lipps, 1976; Moe and Delaca, 1976) and in the South Orkney Islands as well (White and Robins, 1972). Between about 80 and 110 m a soft-bottom epi-infau- nal community of the type discussed by Gallardo (1987) occurred. By wet weight (Figure 4) the dominant compo- nents of this community were sponges (31.4%); echino- derms, chiefl y holothurians (24.6%); ascidians (18.7%); 15_Winston_pg205-222_Poles.indd 21015_Winston_pg205-222_Poles.indd 210 11/17/08 9:22:02 AM11/17/08 9:22:02 AM SYSTEMATICS AND BIOLOGY OF ANTARCTIC BRYOZOANS 211 bryozoans (14.6%) and coelenterates, mostly gorgoni- ans, especially Ophidogorgia paradoxa and Thouarella spp.; and hydroids (10.0%) (Winston and Heimberg, 1988). Over 30 species of bryozoans occurred. The fi ve most abundant species (Carbasea ovoidea, Nematofl us- tra fl agellata, Austrofl ustra vulgaris, Kymella polaris, and Himantozoum antarcticum) all have fl exible, foliaceous, lightly calcifi ed colonies (e.g., Figure 2a,c). Therefore, in terms of volume or area of sea bottom covered, bryozo- ans are an even more important component of the com- munity than would be indicated by biomass alone. Al- though not the highest bryozoan biomass reported from Antarctic benthic communities (Starmans et al., 1999, re- ported 22% bryozoan biomass in Amundsen and Belling- shausen seas), the biomass of bryozoans is typical of the multilayered, microhabitat-rich, fi lter-feeding community reported from many areas of the Antarctic shelf from the Ross Sea (e.g., Bullivant, 1959:488, fi g. 3; Dayton et al., 1974; Dayton, 1990), to King George Island (e.g., Rauschert, 1991:67, fi g. 23), to Signy Island (Barnes, 1995b), to the Weddell Sea (Starmans et al., 1999). Large motile invertebrates found in this habitat in- cluded the isopods Glyptonotus antarcticus and Serolis sp., the echinoid Sterechinus neumayeri, and various aster- oids, e.g., Odontaster validus and Labidaster annulatus. Demersal fi sh species included nototheniids (Notothenia gibberifrons, N. larseni, N. nudifrons, and N. coriiceps) and the icefi sh Chaenocephalus aceratus. Growth and Longevity of Most Abundant Species Life history characteristics of Antarctic bryozoans are amenable to analysis for the following three reasons: (1) Growth is in discrete modules (zooids), making incre- ments of growth easy to measure. (2) Colony fronds of perennial Antarctic species, like those of some temperate species, show yearly growth checks where growth stops during winter months, making it possible to determine the age of fronds as well as their yearly growth rate by back measurement (Stebbing, 1971b; Winston, 1983; P?tzold et al., 1987; Barnes, 1995c; Brey et al., 1998). (3) Embryos, usually brooded in zooid cavities or in ovicells, are easily detectable in living colonies, making it possible to quantify reproductive effort. Carbasea ovoidea was the most abundant bryozoan species in the Low Island community, often comprising more than half the bryozoan biomass in a trawl sample. The delicate, tan, unilaminar fronds of Carbasea colonies showed no growth checks, indicating that all the growth of a particular frond took place during one season, although basal rhizoids and stolons (of this and the other species) might be perennial, producing new fronds yearly. The mean increase in height for Carbasea fronds was 8.6 cm per year. In the other dominant species, both colonies and fronds were perennial. The narrow, curling fronds of Himantzoum could not be accurately measured and so were excluded FIGURE 4. Biomass of organisms from three trawl collections at 100 m, Low Island, Antarctic Peninsula. 15_Winston_pg205-222_Poles.indd 21115_Winston_pg205-222_Poles.indd 211 11/17/08 9:22:03 AM11/17/08 9:22:03 AM 212 SMITHSONIAN AT THE POLES / WINSTON from this part of the study. The other three perennial species grew much more slowly than Carbasea. Figure 5 compares growth in height for individual colonies of all three species as determined by back measurement. Two of them, Austro- fl ustra vulgaris and Nematofl ustra fl agellata are related to Carbasea (family Flustridae). Nematofl ustra fronds showed a mean increase in height of only 0.92 cm per year, whereas Austrofl ustra fronds increased 1.2 cm per year. The ascoph- oran cheilostome Kymella polaris is not closely related to the other species but grew at a similar rate. On average, Kymella fronds increased 1.3 cm in height per year. The oldest fronds of Kymella were at least six years in age, and those of Austrofl ustra and Nematofl ustra were seven years. As none were complete colonies, the genetic individuals they represented may have persisted much lon- ger. The life spans of these species appear somewhat shorter than the 10? 50H11001 year life spans of some of the rigid erect bryozoan species found in deeper Antarctic shelf habitats (Winston, 1983; Brey et al., 1998). However, both growth rates and life spans for both Austrofl ustra and Nemato- fl ustra were in the same range as those of temperate pe- rennials like Flustra foliacea (12 years) (Stebbing, 1971b). Barnes (1995c) also studied growth of Nematofl ustra in shallow water at Signy Island. He found an average yearly increase in height of 0.7 cm and estimated that the oldest colonies at that location were 26 years old. Reproduction Three methods of protecting developing embryos were represented among the dominant species. Austrofl us- tra vulgaris and Nematofl ustra fl agellata brood eggs inter- nally in zooid body cavities, while Kymella polaris broods eggs in ovicells of maternal zooids. In contrast, Carbasea ovoidea broods developing embryos externally in embryo sacs attached in clusters of four or fi ve to the orifi ce of each fertile zooid. The reproductive pattern of Carbasea also contrasted with that of the other common species. Reproductive ef- fort in Carbasea was very high (averaging 2953 embryos per colony) at the time of our fi rst census in late austral summer (1 March) and may have peaked earlier in the summer. By the next sampling period, reproduction had ceased entirely, although zooids, like those of other spe- cies, continued to contain actively feeding polypides throughout the study period. The mean number of embryos per colony was lower in the other species studied: Nematofl ustra, Austrofl ustra, FIGURE 5. Annual growth of individual colonies of three species of Low Island bryozoans determined by back measurement. 15_Winston_pg205-222_Poles.indd 21215_Winston_pg205-222_Poles.indd 212 11/17/08 9:22:04 AM11/17/08 9:22:04 AM SYSTEMATICS AND BIOLOGY OF ANTARCTIC BRYOZOANS 213 Himantozoum, and Kymella, averaging a few hundred per colony at any one census (Table 1). In these species, sexual reproduction was still occurring at the last date sampled. For Kymella and Himantozoum, the two species in which embryos were brooded in ovicells, we could determine the percentage of ovicells brooding embryos versus empty ovi- cells over the course of the season (Table 2). The percent- age of empty ovicells gradually increased until by 20 April, 72% of all Kymella ovicells and 95% of all Himantozoum ovicells were empty, indicating that the end of the repro- ductive season was approaching for at least these two spe- cies. In Nematofl ustra the number of embryos fl uctuated slightly from census to census but showed no signifi cant decline as of 20 April. Austrofl ustra vulgaris colonies con- tained mature embryos at the 7 March census (mean of 258 per colony), while colonies collected on 20 April had an average of 227 embryos per colony. Although there was no way for us to tell how long the embryos present in late April would be brooded, it may well be as later workers (e.g., Barnes and Clarke, 1995; Bowden, 2005; Knox, 2007) have suggested, that ?winter? may not be as long for the benthos as predicted on the basis of light and phytoplankton abundance. Partial Mortality and Predation Like most clonal organisms, bryozoans have retained extensive powers of regeneration and can tolerate a high degree of injury or death of portions of the colony without death of the entire colony. Such partial mortality may be caused by physical disturbance of the environment or by predators or grazers. A few invertebrates, including some pycnogonids and nudibranchs, are specialized as single- zooid predators of bryozoans. These animals pierce a zo- oid with proboscis or radula and suck out body fl uids and tissues, leaving an empty or broken zooid behind. Single or small patches of empty zooids were probably the result of such predators. The grazing or browsing activities of fi sh, echinoids, and mollusks leave larger scrapes, rips, and bites on colony fronds. We examined colonies for injuries of the different types and noted where they occurred on the fronds. Table 3 shows that all species sustained a consid- erable amount of damage. Carbasea colonies showed the least amount of injury to growing tips (the most delicate and accessible portion of the frond). This is most likely due to their much higher growth rate. Evidence from studies of gut contents of associated macrobenthic organisms also suggested that most of the injuries observed were not due to feeding by specialized bryozoan predators. A search of the literature revealed that small quantities (from less than 1% to about 3%) of bryozoans had been found in gut contents of several Antarctic fi sh and echinoderms (Dayton et al., 1974; Dearborn, 1977). We examined gut contents of a number of invertebrates from Low Island trawls (including poly- chaetes, echinoderms, and crustaceans) to learn whether any of them were feeding on bryozoans. Two Low Island invertebrates, the echinoid Sterechinus neumayeri and the isopod Glyptonotus antarcticus, did contain bryozoan fragments. But our results, like those of later workers, in- dicated that gut contents of invertebrate carnivores and scavengers, like those of the demersal fi sh, consisted pri- marily of small motile invertebrates: amphipods, isopods, polychaetes, and mollusks (Schwartzbach, 1988; Ekau and Gutt, 1991: McClintock, 1994). Another source of damage to colonies is caused by fouling of colonies by other organisms. Table 4 shows the most common organisms found attached to frontal surfaces of colonies of the fi ve most abundant Low Is- land species. Some of them, such as the stalked barnacles found in branch bifurcations or the colonies of the bryo- zoan Beania livingstonei, which attach loosely to the host colony, may benefi t the host, augmenting colony water currents by their own feeding activities. Others, such as TABLE 1. Mean number of embryos produced per colony for most abundant Low Island species. Mean Number Total number number of Species of embryos per colony colonies Carbasea 15,355 1536 10 Austrofl ustra 5920 321 20 Himantozoum 6326 452 14 Kymella 3698 247 15 Nematofl ustra 19,277 741 26 TABLE 2. Percent of ovicells empty for two Low Island species at three census periods. Species 1 March 2 April 20 April Kymella 19% 40% 75% Himantozoum 46% 62% 95% 15_Winston_pg205-222_Poles.indd 21315_Winston_pg205-222_Poles.indd 213 11/17/08 9:22:05 AM11/17/08 9:22:05 AM 214 SMITHSONIAN AT THE POLES / WINSTON encrusting bryozoans like Ellisina and Harpecia, disable the host zooids they overgrow. Table 5 shows the diversity and density of fouling on Low Island species compared with that on NE Atlantic Flustra foliacea, as studied by Stebbing (1971a). The overall number of taxa and num- ber of epizoans per colony was lower for all the Low Is- land species studied than for Flustra. However, when the number of epizoans per square centimeter was calculated, two species, Austrofl ustra (1.10/cm 2 ) and Kymella (1.2/ cm 2 ), were in the same range as Flustra foliacea (1.0/cm 2 ). Barnes (1994) studied the epibiota of two erect species, Alleofl ustra tenuis and Nematofl ustra fl agellata, from shallow (36? 40 m) and deeper (150 m) habitats at Signy Island. The frontal surface of Nematofl ustra showed fewer colonizers than the abfrontal surface, and for both species the amount of fouling decreased to almost zero at 150 m. The number of taxa encrusting both species was also low (median H11005 3.0 for Alleofl ustra and 2.0 for Nematofl us- tra). Our methods were somewhat different, but it appears that overall diversity of epibiotic taxa was higher at Low Island, but the diversity of epibiotic bryozoan taxa was much higher at Signy Island. Food Sources Gut contents of the bryozoans studied are summarized in Table 6. Each autozooid polypide has a mouth at the base of the lophophore funnel. Mouth size is slightly vari- able, as the mouth expands and contracts slightly with par- ticle swallowing, but it is closely correlated with zooid size (Winston, 1977). These Antarctic species had large mouths compared to species from warmer water but, somewhat sur- prisingly, were still feeding primarily on very small plank- ton cells, mostly tiny diatoms and dark, rough-walled cysts less than 20 H9262m in size, probably resting stages of either choanofl agellates (Marchant, 1985; Marchant and McEl- downy, 1986) or diatoms (Bodungen et al., 1986). Most of the phytoplankton component of their diet was thus within the nanoplankton, a size range which has been shown to account for much of the primary productivity in some areas of Antarctic seas (Bracher, 1999; Knox, 2007). The Brans- fi eld Strait area is an important breeding ground for krill, which feed on larger plankton. Nanoplankton and pico- plankton populations increase as microplankton blooms diminish (Varela et al., 2002). Some studies have found TABLE 3. Mean number of injuries per colony and percent of branch tips injured. Number of Number of Number of Number of rips torn injuries to bites to injuries to Percentage of in branches branch tip branch edges center of a Number of branch tips Species (growing edge) (sides of branches) branch empty zooids injured Carbasea 1.7 3.4 1.1 1.4 130 4.4% Nematofl ustra 4.0 3.5 5.0 7.2 41 23.5% Austrofl ustra 1.4 2.8 3.0 3.8 81 52% Kymella 1.2 4.1 0.5 0.2 50 53% TABLE 4. Dominant epizoans attached to dominant Low Island bryozoan species; a ?H11001? indicates their presence on a particular bryo- zoan species. Epizoan organisms Carbasea Nematofl ustra Flustra Himantozoum Kymella Foraminiferans H11001 H11001 H11001 Diatoms H11001 Hydroids H11001 H11001 Stalked barnacles H11001 H11001 Beania livingstonei H11001 H11001 Harpecia spinosissima H11001 Ellisina antarctica H11001 H11001 Osthimosia sp. H11001 Cyclostome bryozoans H11001 15_Winston_pg205-222_Poles.indd 21415_Winston_pg205-222_Poles.indd 214 11/17/08 9:22:05 AM11/17/08 9:22:05 AM SYSTEMATICS AND BIOLOGY OF ANTARCTIC BRYOZOANS 215 nanoplankton making up 83% of phytoplankton carbon in February and March (Kang and Lee, 1995). Sediments in the peninsula are also rich in organic matter (Bodungen et al., 1989). Benthic microalgae (Vincent, 1988; Gilbert, 1991) may also be important in areas reached by light. The bryozoans studied also ingested large amounts of ?brown particulate material.? This material contained chlorophyll and may have been derived from the fecal pel- lets of zooplankton or those of other benthos. Fecal mate- rial could also have derived from benthos (N?thig and von Bodungen, 1989; Tati?n et al., 2004). Feeding on dead or- ganisms or fecal material has been shown to occur in other Antarctic animals and may aid them in surviving seasons of low food supply. Sediment particles may indicate the importance of an advected food supply, as Gutt (2007) has speculated, which might explain the sediment grains found in bryozoan gut contents. Some food resource besides phytoplankton seems likely to be part of Antarctic bryozoan life histories. Pol- ypides of colonies of all species we observed were still ac- tively feeding at the end of our stay in late April. This is in keeping with the observations made by Barnes and Clarke (1994), who monitored colony activity every month for a two-year period at Signy Island. Most bryozoans they observed stopped feeding for only a two- to three-month period in mid? austral winter. Habitat and Ecosystem Role The primary role of bryozoans in the Low Island eco- system appeared to be that of habitat and foraging ground for demersal fi sh and motile invertebrates. To assess their importance in that regard, we selected three large clusters of Carbasea ovoidea from a single trawl and immediately immersed them in large jars of seawater formalin to kill and preserve their inhabitants for later analysis. Table 7 shows the results. When living, these clumps contained about 0.09 m 3 of habitat space (in overall volume, not counting areas of each frond) and held almost 500 indi- vidual invertebrates. Most were arthropods (81.6%), of which amphipods comprised the majority: 345 individu- als (69.6%). Nematodes were the next largest group pres- ent, with 54 individuals (10.8%). Other motile and sessile groups made up the remainder of the inhabitants. Most of them fi gure in the diets of demersal fi shes and benthic invertebrates of the Antarctic Peninsula region (Targett, 1981; Daniels, 1982). Figure 6 is a diagrammatic representation of a Low Island food web, placing the bryozoans within it as part of the link to the benthos. The major role of bryozoans within the Low Island ecosystem is as a three-dimensional, layered habitat and shelter for the small invertebrates on which larger demersal fi sh and benthic invertebrates feed TABLE 5. Diversity and density of epizoans fouling Low Island bryozoans compared with fouling on northeast Atlantic Flustra foliacea. Species Number of epizoic taxa Mean number of epizoans/colony Mean number/cm 2 of colony surface Carbasea ovoidea 9 28 0.56 Nematofl ustra fl agellata 13 14 0.24 Austrofl ustra vulgaris 12 36 1.10 Himantozoum antarcticum 9 9 ? Kymella polaris 9 28 1.2 Flustra foliacea 42 558 1.0 TABLE 6. Gut contents of Low Island bryozoans. Species Mean mouth size (H9262m) Dominant particle types a Particle size range (H9262m) Carbasea ovoidea 59 BPM, diatoms, cysts 3? 66 Austrofl ustra vulgaris 96 BPM, cysts 45 Kymella polaris 94 BPM, sediment grains 6? 60 Nematofl ustra fl agellata 87 Cysts, diatoms 9? 69 Beania livingstonei 114 BPM, diatoms, cysts, sediment grains 3? 93 a BPM H11005 brown particulate material. 15_Winston_pg205-222_Poles.indd 21515_Winston_pg205-222_Poles.indd 215 11/17/08 9:22:05 AM11/17/08 9:22:05 AM 216 SMITHSONIAN AT THE POLES / WINSTON and a part of the epibenthic nursery ground for juveniles of demersal fi sh and motile invertebrates. Along with other sessile epifauna, they may also provide large epibi- otic fi lter-feeding invertebrates with a higher perch and better access to food (Gutt and Shickan, 1998). The bryo- zoans themselves are suspension-feeding consumers of mi- croplankton and nanoplankton, as well as fecal material or detritus supplied from above their colonies. They may also play a minor role as a food source for single-zooid predators and a few fi sh and invertebrates. BEHAVIOR OF ANTARCTIC BRYOZOANS Morphological studies of avicularia and vibracula (Winston, 1984) and behavioral studies carried out with living colonies (Winston, 1991) have shown how different polymorphs, including those of Antarctic species, function within colonies. These functions include sweeping debris from zooids and colonies, protecting the colony from trespassers, and capturing motile organisms and possible predators. NEMATOFLUSTRA? VIBRACULA Distal to each autozooid of Nematofl ustra fl agellata is a vibraculum zooid with a long, curved, bristle-like man- dible (Figure 7a). Mechanical stimulation or the vibration of a small organism, such as a pycnogonid or polychaete, triggered a wave of vibracular movement over a part or all of the colony surface. The circumrotary reversal of the vi- bracula mandibles was sequentially synchronized at fi rst, traveling proximally from the branch tip, but later waves became less synchronous. The waves of moving setae ef- fectively carried organisms or debris from the branches (Figure 7b) (Winston, 1991). BEANIA? BIRD?S HEAD AVICULARIA Unlike sessile avicularia and vibracula, the peduncu- late (bird?s head) avicularia found in the cellularine group of bryozoans close and open mandibles frequently, even when undisturbed, showing a species-specifi c pattern of ongoing activity. For example, in Beania livingstonei (Fig- ure 7c) the avicularium bends forward on its peduncle, then snaps back into an upright position, while the man- dible closes. Once the avicularium is upright, the mandible slowly reopens. Although the avicularian movements did not increase in the presence of trespassers, their activity was still effective as organisms caught by one avicularium were usually captured by the mandibles of several more in their struggles to free themselves (e.g., Figure 7d). Some were eventually able to pull free, detaching the avicularia from their peduncles in the process (Winston, 1991). CAMPTOPLITES? LONG-STALKED BIRD?S HEAD AVICULARIA The long-stalked avicularia of Camptoplites species (Figure 7e) show an even more complex pattern. The long, slender peduncles of these avicularia sway slowly back and forth across the frontal surface of colony branches. As they sway, the mandibles of the avicularia snap open and TABLE 7. Animals inhabiting three Carbasea ovoidea colonies collected at Low Island. Group Number of specimens Amphipods 345 Nematodes 54 Isopods 41 Pycnogonids 19 Ascidians 17 Bivalves 12 Holothurians 4 Sponges 2 Sipunculans 1 Asteroids 1 Total 496 FIGURE 6. Low Island food web. Diagrammatic representation of links between benthic and pelagic communities. 15_Winston_pg205-222_Poles.indd 21615_Winston_pg205-222_Poles.indd 216 11/17/08 9:22:06 AM11/17/08 9:22:06 AM SYSTEMATICS AND BIOLOGY OF ANTARCTIC BRYOZOANS 217 FIGURE 7. Avicularia and vibracula of some Antarctic bryozoan species studied. (a) Living branch of Nematofl ustra with vibracula in un- disturbed position. (b) Vibracula-sweeping polychaete from colony surface (still photo from video). (c) Beania livingstonei. Arrow points to one of the bird?s head avicularia. (d) Nematode caught by several avicularia (still photo from video). (e) Long-stalked bird?s head avicularia of Camptoplites colony. (f) Worm speared by one of the sharp-pointed mandibles of Micropora brevissima. Tiny arrow points to mandible inside worm?s translucent body. 15_Winston_pg205-222_Poles.indd 21715_Winston_pg205-222_Poles.indd 217 11/17/08 9:22:06 AM11/17/08 9:22:06 AM 218 SMITHSONIAN AT THE POLES / WINSTON shut. They show no increase in activity when a trespass- ing organism touches the branches, but when a mandible intercepts an object soft and narrow enough to grasp (such as a polychaete chaeta or arthropod appendage), it snaps shut upon it. The swaying activity then carries the organ- ism toward the edge of the colony (Winston, 1991). MICROPORA? SESSILE AVICULARIA Micropora brevissima (Figure 7f) has sessile avicularia with sharply pointed mandibles. In avicularia of this type, the bodies of the avicularia are fi xed on the colony, and their mandibles close rarely, remaining in an open position even when zooid opercula are shut. The mandible shuts in response to physical stimuli, probing, jarring, or vibration of palate or mandible. Despite their small size, such avicu- laria are able to capture and hold relatively large trespass- ers by their appendages, e.g., cirri of polychaetes or legs of amphipods or pycnogonids. During observations of living colonies of this species carried out at Palmer Station, avic- ularia of Micropora captured several annelids and held them despite their much greater size (Winston, 1991). CONCLUSIONS Antarctic bryozoans are taxonomically rich and, like many other Antarctic organisms, show a high degree of endemism. Preliminary analysis of USARP bryozoan col- lections from the Ross Sea and Antarctic Peninsula yielded 222 species from all three marine orders: fi ve ctenostomes, 28 cyclostomes, and 189 cheilostomes. Hayward (1995) studied Antarctic cheilostomes, primarily from British Antarctic research programs but also including some in- formation from USARP collections. He listed 264 spe- cies of cheilostomes, of which 215 (81%) were endemic. Moyano (2005) took a different approach in summariz- ing taxonomic research on Antarctic bryozoans through 2004. He included taxa from all three orders, but some of his totals included subantarctic species. His list totaled 315 species, 250 (79%) of which were endemic. Antarctic bryozoans are abundant. It is hard to fi nd a published underwater benthic photograph from the Ant- arctic in which a bryozoan colony cannot be seen, and in many areas, their skeletons, along with sponge spicules and glacial pebbles, make up a signifi cant component of the sediment. As shown in results from work at Low Island and elsewhere, bryozoans play a signifi cant role in benthic ecosystems, providing habitat and foraging ground as well as a minor food resource for fi sh and motile invertebrates. Many of the erect, branching, and foliose bryozoans which form such three-dimensional habitats are perennial, with life spans lasting several years, perhaps several decades. Their growth rates are slow and their fecundity low. Car- basea ovoidea, the exception to this pattern at Low Is- land, is a species whose northernmost range extends into the subantarctic and probably reaches the southernmost extent of its range somewhere along the peninsula (Hay- ward, 1995). Most of the Antarctic fi sh and many other benthic in- vertebrates that feed among the bryozoans are similarly slow growing and long-lived. Their survival has depended on long-term stability of their environment. Two factors, trawling (illegal fi shing), which has already decimated stocks of the most desirable fi sh species, and global cli- mate change, could have severe consequences for such communities. The impact of climate change will differ depending on the temperature scenario. As pointed out by Thatje et al. (2005), the lowering of temperatures, leading to a gla- cial period, could almost completely eliminate the benthic fauna of the Antarctic shelf and slope. Increasing tempera- tures (as seen today, especially in the West Antarctic) will probably lead to enhanced diversity in a few habitats (e.g., the intertidal zone of the peninsula) but also to a change in present-day communities due to human impact and tem- perature stress. Human impact, including increased travel, both scientifi c and touristic, will increase the likelihood of ship fouling (Lewis et al., 2005; Barnes and Conlan, 2007). In addition, an increase in growth rate with higher temperatures has already been detected in one bryozoan, Cellarinella nutti (Barnes et al., 2006). As Collie et al. (2000) noted in their analysis of the impacts of fi shing on shelf benthos, there are large gaps in our knowledge of the effects of trawling on three-dimen- sional epifaunal communities. Reduction in habitat com- plexity caused by intensive bottom fi shing may have long- term negative consequences for fi sh communities (loss of nest sites and nursery and breeding grounds). Although benthic habitats with a high degree of struc- tural heterogeneity occur in areas besides Antarctica, such as New Zealand (Bradstock and Gordon, 1983; Batson and Probert, 2000) and Helgoland (de Kluijver, 1991), they are rare. In only one (Tasman Bay, New Zealand) have erect bryozoan communities been protected in an at- tempt to restore a fi shery (Bradstock and Gordon, 1983). Finally, despite all the biological work that has oc- curred in the Antarctic since the 1980s, we still have a very poor idea of the link between what happens in pe- lagic communities and the benthos. A solid understanding 15_Winston_pg205-222_Poles.indd 21815_Winston_pg205-222_Poles.indd 218 11/17/08 9:22:15 AM11/17/08 9:22:15 AM SYSTEMATICS AND BIOLOGY OF ANTARCTIC BRYOZOANS 219 of benthic-pelagic coupling, factoring in life histories of structural epibenthos and their epibionts as well as resting stages of phytoplankton, importance of fecal pellets, and the role of benthic meiofauna in food chains and shelf eco- systems, has hardly begun (Marcus and Boero, 1998). ACKNOWLEDGMENTS Thanks go to the staff members of SOSC and staff and volunteers at American Museum of Natural History and Virginia Museum of Natural History, who sorted thousands of specimens, and to my colleagues Peter J. Hayward, Uni- versity of Wales, Swansea, for making several trips to the United States to work on the collections with me, sorting, identifying, and verifying preliminary identifi cation, and Beverly Heimberg, American Museum of Natural History, for her work with me at Palmer Station and American Mu- seum of Natural History. Thanks also go to the Smithsonian Institution for the loan of the specimens and several SOSC and Invertebrate Biology contracts that helped support the sorting, identifi cation, and cataloging of results and to the National Science Foundation for NSF grant DPP-8318457 (1984), which supported the work on ecology and behav- ior of living Antarctic bryozoans. Thanks to support staff and other scientists at the station in late summer and fall of 1985 and to the crew of the Polar Duke for their logistical support and entertainment. LITERATURE CITED Arnaud, P. M. 1974. Contribution ? la Bionomie Marine Benthic des Regions Antarctiques et Subantarctiques. 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Hayward. 1994. ?Bryozoa of the U.S. Antarctic Research Program: A Preliminary Report.? In Biology and Paleo- biology of Bryozoans, ed. P. J. Hayward, J. S. Ryland, and P. D. Taylor, pp. 205? 210. Fredensborg, Denmark: Olsen & Olsen. Winston, J. E., and B. F. Heimberg. 1988. The Role of Bryozoans in the Benthic Community at Low Island, Antarctica. Antarctic Journal of the United States, 21: 188? 189. 15_Winston_pg205-222_Poles.indd 22115_Winston_pg205-222_Poles.indd 221 11/17/08 9:22:16 AM11/17/08 9:22:16 AM 15_Winston_pg205-222_Poles.indd 22215_Winston_pg205-222_Poles.indd 222 11/17/08 9:22:16 AM11/17/08 9:22:16 AM ABSTRACT. Interdisciplinary studies of narwhal cranial and tooth anatomy are combined with Inuit traditional knowledge to render a more complete description of tooth- related structures and to propose a new hypothesis for tusk function in the adult male. Gross anatomy fi ndings from computed tomography (CT) and magnetic resonance (MR) imag- ing and dissections of an adult male and female and one fetus, four to six months in devel- opment, were documented. Computed tomography scans rendered images of the tusks and vestigial teeth and their shared sources of innervation at the base of the tusks. Paired and asymmetrical tusks and vestigial teeth were observed in all three samples, and their relative positions reversed during development. Vestigial teeth shifted anteriorly during growth, and the developing tusks moved posteriorly as they developed. Examination of tusk micro- anatomy revealed the presence of a dentinal tubule network with lumena approximately 2 micrometers in diameter and 10? 20 micrometers apart over the pulpal and erupted tusk surfaces. Orifi ces were present on the cementum surface indicating direct communication and sensory capability from the environment to the inner pulpal wall. Flexural strength of 95 MPa at mid tusk and 165 MPa at the base indicated resistance to high fl exural stresses. Inuit knowledge describes a tusk with remarkable and combined strength and fl exibility. Elder observations of anatomy are described by variable phenotypes and classifi ed by skin coloration, sex, and tusk expression. Behavioral observations of males leading seasonal migration groups, nonaggressive tusk encounters, and frequent sightings of smaller groups separated by sex add to the discussion of tusk function. INTRODUCTION The narwhal is unique among toothed marine mammals and exhibits unusual features, which are described in the literature (Figure 1). A single 2? 3 m tusk is characteristic of adult males (Tomlin, 1967). Tusks are horizontally imbedded in Martin T. Nweeia, Harvard University, School of Dental Medicine, Boston, MA, USA. Cornelius Nutarak and David Angnatsiak, Community of Mittimatilik, Nunavut, Canada. Frederick C. Eichmiller, Delta Dental of Wisconsin, Stevens Point, WI, USA. Naomi Eidelman, Anthony A. Giuseppetti, and Janet Quinn, ADAF Paffen- barger Research Center, National Institute of Stan- dards and Technology, Gaithersburg, MD, USA. James G. Mead and Charles Potter, Smithsonian Institution, Division of Mammals, Department of Zoology, Washington, DC, USA. Kaviqanguak K?issuk and Rasmus Avike, Community of Qaa- naaq, Greenland. Peter V. Hauschka, Smithsonian Institution, Division of Mammals, Department of Zoology, Washington, DC, USA. Ethan M. Tyler, National Institutes of Health, Clinical Center, Bethesda, MD, USA. Jack R. Orr, Fisheries and Oceans Canada, Arctic Research Division, Winni- peg, MB, Canada. Pavia Nielsen, Community of Uummannaq, Greenland. Corresponding author: M. T. Nweeia (martin_nweeia@hsdm.harvard. edu). Accepted 19 May 2008. Considerations of Anatomy, Morphology, Evolution, and Function for Narwhal Dentition Martin T. Nweeia, Frederick C. Eichmiller, Cornelius Nutarak, Naomi Eidelman, Anthony A. Giuseppetti, Janet Quinn, James G. Mead, Kaviqanguak K?issuk, Peter V. Hauschka, Ethan M. Tyler, Charles Potter, Jack R. Orr, Rasmus Avike, Pavia Nielsen, and David Angnatsiak 16_Nweela_pg223-240_Poles.indd 22316_Nweela_pg223-240_Poles.indd 223 11/18/08 9:12:43 AM11/18/08 9:12:43 AM 224 SMITHSONIAN AT THE POLES / NWEEIA ET AL. the upper jaw and erupt through the left side of the max- illary bone, while the smaller tusk on the right side, usu- ally not longer than 30 cm, remains embedded in the bone. Narwhal thus exhibit an extreme form of dental asymmetry (Hay and Mansfi eld, 1989; Harrison and King, 1965). One and a half percent of narwhal exhibit double tusks. Such expression is marked by a right tusk that is slightly shorter than the left (Fraser, 1938) and has the same left-handed helix surface morphology (Clark, 1871; Gervais, 1873; Thompson, 1952). An expected tusk antemere would be equal in size and have a mirror-imaged morphology. Thus, narwhal dental asymmetry is uncharacteristic in size and shape. Only in rare instances, such as the fossil record of Odobenoceptops peruvianus, a walrus-like cetacean hypo- thesized to be in the Delphinoidae family and possibly re- lated to Monodontidae, does such tusk asymmetry exist (de Muizon et al., 1999; de Muizon and Domning, 2002). Thus, narwhal dental asymmetry is uncharacteristic in size and shape. Erupted tusks pierce through the lip, while the embed- ded tusk remains in bone. Tusk surface morphology is dis- tinguished by a characteristic left-handed helix (Kingsley and Ramsay, 1988; Brear et al., 1990) rarely seen in other tusked mammals, with the exception of unusual examples like elephants that undergo trauma shortly after birth and develop spiraled tusks (Busch, 1890; Colyer, 1915; Goethe, 1949). Most male narwhal have a tusk, while only 15% of females have a tusk (Roberge and Dunn, 1990). When exhibited, female tusks are shorter and narrower than male tusks (Clark, 1871; Pedersen, 1931). Narwhal tusk expression is thus an unusual example of sexual di- morphism in mammalian teeth. Comparative fi ndings are limited for other beaked whales that exhibit sexually dimorphic teeth (Heyning, 1984; Mead, 1989). Fetal nar- whal develop six pairs of maxillary teeth and two pairs of mandibular teeth. Only two pairs of upper teeth form in the adult narwhal, the second pair being vestigial and serv- ing no known function. FIGURE 1. Male narwhal whales with their characteristic tusks. 16_Nweela_pg223-240_Poles.indd 22416_Nweela_pg223-240_Poles.indd 224 11/18/08 9:12:43 AM11/18/08 9:12:43 AM CONSIDERATIONS OF NARWHAL DENTITION 225 Many theories have been hypothesized to explain the purpose and function of the erupted tusk. Proposed ex- planations include a weapon of aggression between males (Brown, 1868; Beddard, 1900; Lowe, 1906; Geist et al., 1960), a secondary sexual characteristic to establish social hierarchy among males (Scoresby, 1820; Hartwig,1874; Mansfi eld et al., 1975; Silverman and Dunbar, 1980), an instrument for breaking ice (Scoresby, 1820; Tomlin, 1967), a spear for hunting (Vibe, 1950; Harrison and King, 1965; Bruemmer, 1993:64; Ellis, 1980), a breathing organ, a thermal regulator, a swimming rudder (Kingsley and Ramsay, 1988), a tool for digging (Freuchen, 1935; Pederson, 1960; Newman, 1971), and an acoustic organ or sound probe (Best, 1981; Reeves and Mitchell, 1981). Examination of tusk anatomy, histology, and biome- chanics combined with traditional knowledge of Inuit el- ders and hunters has revealed features that support a new sensory hypothesis for tusk function (Nweeia et al., 2005). Narwhal tusk reaction and response to varying salinity gra- dients introduced during fi eld experiments support this. GROSS CRANIAL AND TOOTH ANATOMY Three narwhal head samples, obtained during legal Inuit harvests in 2003 and 2005 were examined by com- puterized axial tomography and magnetic resonance im- aging and then dissected. They included one adult male, one adult female, and one fetal specimen between four and six months in its development. The department of ra- diology at Johns Hopkins Hospital conducted computed tomography (CT) scans on all three specimens using a Sie- mens Medical Solutions SOMATOM Sensation Cardiac 64. The scanner generated 0.5-mm-thick slices on each of the three specimens. Original data from these scans has been archived at the Smithsonian Institution. Material- ize Mimics 8.0 and Discreet 3D Studio Max 7 was used to create digital 3-D models of narwhal dental anatomy. Magnetic resonance imaging (MRI) was also used to in- vestigate and visualize narwhal dental anatomy. Scientists at the National Institutes of Health MRI Research Facil- ity conducted MRI on the thawed narwhal heads. Data from MRI assisted verifi cation of known cranial anatomy and enabled examination of tooth vasculature. The nar- whal heads were dissected at the Osteo-Prep Laboratory at the Smithsonian Institution, and digital photographs were taken to record anatomical landmarks and features of gross anatomy. The skull of the fetus was 6.23 cm in length and 4.96 cm in width at the most distal points on its frontal bones (Figure 2). Computed tomography data were col- lected for the entire body of the fetus, though only the head was investigated for the purpose of this research. The total length of the specimen was 31.68 cm. Calcifi - cation of major bones of the skull was incomplete in the fetal narwhal specimen. The top of the skull was smoothly rounded, with a large anterior fontanelle, and there were lateral vacuities where ossifi cation was incomplete. Most of the main membrane bones were present at this stage. The nasal bones were small elliptical bones positioned on the summit of the head dorsal to the tectum nasi; they did not make medial contact. The premaxillae were long, narrow shafts of bone medial to maxillae. The maxillae were large bones that were excavated anteroventrally to form two conspicuous pairs of tooth sockets, or alveoli. The maxillae overlapped the frontal bones. Pre- and post- orbital processes were well developed. The parietal bones were small lateral bones that made contact with frontal FIGURE 2. A drawing made from CT scans of the skull and teeth of a narwhal fetus showing the positions of the teeth. 16_Nweela_pg223-240_Poles.indd 22516_Nweela_pg223-240_Poles.indd 225 11/18/08 9:12:47 AM11/18/08 9:12:47 AM 226 SMITHSONIAN AT THE POLES / NWEEIA ET AL. bones on their anterior borders. As some of the bone was very thin and articulated with cartilage, digital models of the specimen had some minor artifacts. Two pairs of teeth were evident in the upper jaw of the fetal narwhal specimen. Future tusks and vestigial teeth were located in their respective sockets in the maxillary bones. The future tusks were located anteromedially to the vestigial pair of teeth. These moved in a posterior direc- tion during development, forming the two large tusks in the adult narwhal (Figure 3). In the male, the left future tusk usually becomes the erupted tusk, and the right tusk typically remains embedded in the maxilla. The future tusks exhibited asymmetry, with the right being 0.44 cm in length and 0.38 cm in width at its widest point and the left being 0.36 cm length and 0.36 cm in width at its wid- est point. The vestigial pair of teeth also exhibited asymmetry. The right vestigial tooth was 0.26 cm in length and 0.25 cm in width at its widest point, and the left vestigial tooth was 0.32 cm in length and 0.25 cm in width at its widest point. No teeth were evident in the mandible of this fetus, though dental papillae in the lower jaw have been docu- mented in a report that describes up to six pairs of teeth in the upper jaw and two pairs in the lower jaw (Eales, 1950). The vestigial pair of teeth and their shared blood and nerve supply with the tusks suggest that the narwhal may have exhibited at least two well-developed pairs of teeth at some point in its evolution. The head of the adult female narwhal was 55.82 cm in length and 47.90 cm in width at its base. The skull of the specimen was 53.96 cm in length and 35.08 cm in width at the level of the bases of the embedded tusks. Like most ce- taceans in the family Odontoceti, the skull of the narwhal was asymmetrical, with bony structures skewed toward the left side of the head. Two pairs of teeth were visible in the upper jaw of the female narwhal specimen (Figures 4 and 5). Both pairs, tusks and vestigial, were found in their respective sockets in the maxillae. The tusks were located posteromedially to the vestigial pair. In the female, the paired tusks typically remain embedded in the maxillae, as was the case for this specimen. The tusks exhibited asymmetry. The right tusk was 17.47 cm in length, 2.39 cm in width at its base, and 0.80 cm in width at its distal end. The left tusk was 18.33 cm in length, 2.06 cm in width at its base, and 1.07 cm in width at its distal end. In rare cases, the left tusk of the female erupts from the maxilla. Sockets for the tusks in the skull begin at the bases of the teeth and terminate in the most distal part of each respective maxillary bone. The vestigial pair of teeth also exhibited asymmetry. The right vestigial tooth was 2.39 cm in length and 0.98 cm in width at its widest point. The left vestigial tooth was 1.90 cm in length and 0.98 cm in width at its widest point. Sockets for the vestigial teeth began near the bases of the tusks and, like the tusks, terminated in the most distal part of each maxillary bone. The right vestigial tooth slightly protruded from the bone. During dissection and prepara- tion, vestigial teeth may be lost because they are not se- curely embedded in the bone. There is limited documenta- tion on the presence and morphology of vestigial teeth. Evidence of developed sockets for the vestigial teeth in the narwhal is signifi cant, as it suggests that this species may have exhibited at least two well-developed pairs of teeth FIGURE 3. A drawing showing the migration and reversal of position of the teeth from fetus to adult that occurs during development. 16_Nweela_pg223-240_Poles.indd 22616_Nweela_pg223-240_Poles.indd 226 11/18/08 9:12:52 AM11/18/08 9:12:52 AM CONSIDERATIONS OF NARWHAL DENTITION 227 at some point in its evolution. The two pairs of teeth also effectively reverse positions during development. On the basis of intracranial dissection of the trigemi- nal or fi fth cranial nerve, the optic nerve branch passed through the superior orbital fi ssure (Figure 6). The maxil- lary branch, a sensory nerve, passed through the foramen rotundum, and the mandibular nerve branch, a motor and sensory nerve, passed through the foramen ovale. The head of the male narwhal was 62.33 cm in length and 49.70 cm in width at its base. The skull of the speci- men was 57.24 cm in length and 35.01 cm in width at the level of the base of the left tusk. Like the female narwhal, the skull of the male narwhal was asymmetrical, and bony structures skewed toward the left side of the head. FIGURE 4. A three dimensional reconstruction made from CT data of the adult female dentition of M. monoceros showing (A) the dor- sal view and (C ) lateral view of the unerupted tusks with (B) detail of left vestigial tooth. FIGURE 5. (A) Lateral view of the complete female head with coro- nal cuts (B) and (C). (B) shows the embedded tusks and vestigial teeth and (C) shows vestigial teeth with vascularized tusk sockets. 16_Nweela_pg223-240_Poles.indd 22716_Nweela_pg223-240_Poles.indd 227 11/18/08 9:13:01 AM11/18/08 9:13:01 AM 228 SMITHSONIAN AT THE POLES / NWEEIA ET AL. Two pairs of teeth were visible in the upper jaw of the male narwhal specimen (Figure 7). The paired tusks were located posteromedially to the vestigial pair of teeth. As is typical with the species, the right tusk remained embedded in the skull, and the left tusk erupted from the maxilla. The right tusk was 20.81 cm in length and 2.51 cm at its base; it was 0.96 cm in width at its distal end. The left tusk was 89.62 cm in length and 4.33 cm in width at its base. It was 4.36 cm in width as it exited the maxilla, and 2.96 cm in width at the termination of CT data at 56.45 cm distal to the maxilla. Bony sockets for the tusks began at the bases of the teeth and terminated in the most distal part of each respective maxillary bone. The vestigial pair of teeth also exhibited asymmetry. Unlike the vestigial teeth of the female specimen, the vestigial teeth of the male narwhal were not embedded in bone; they were suspended in the tissue lateral to the maxillae. The right vestigial tooth was 0.59 cm in length and 0.21 cm in width at its widest point; the left vestigial tooth was 0.67 cm in length and 0.20 cm in width at its widest point. The Inuit classify narwhal with separate names on the basis of skin color and tusk expression. For example, a male with black coloration is eiJ4g6, and a male with white coloration is cfJ4g6. Males with a shorter and wider tusk are called g]Zwg8, and those with a longer, nar- rower tusk and black skin coloration are gZE8i3nw. Sev- eral Inuit from High Arctic communities in northwestern Greenland are able to recognize and differentiate narwhal populations from Canada and Greenland by their body form and their behavior. They describe Canadian narwhal as being narrower through the length of their bodies and more curious and social, while the Greenlandic narwhal FIGURE 6. (A) Three dimensional reconstruction made from CT data showing the posterior view of the adult female M. monoceros skull with the access opening for intracranial dissection, and (B, C) a drawing from photographs taken during dissection showing the nerves and foramen at the cranial base. 16_Nweela_pg223-240_Poles.indd 22816_Nweela_pg223-240_Poles.indd 228 11/18/08 9:13:11 AM11/18/08 9:13:11 AM CONSIDERATIONS OF NARWHAL DENTITION 229 are wider and more bulbous in the anterior two thirds of their bodies and taper at the tail. Their personalities are described as being shyer, and thus they are more elusive. GROSS TUSK MORPHOLOGY A 220-cm-long tusk harvested in 2003 at Pond Inlet, Nunavut, Canada, during the Inuit hunting season was sectioned in the fi eld using a reciprocating saw into trans- verse slices averaging approximately 5 cm in length. Sec- tioning was started approximately 40 cm inside the skull from the point of tusk eruption and as close to the devel- oping base as possible. The sections were immersed in an aqueous solution of 0.2% sodium azide and frozen in indi- vidual serially identifi ed containers. The sections were fi rst evaluated for gross features and dimensions while intact, and selected specimens were further sectioned for more detailed microscopy. Gross section measurements revealed an outer diam- eter tapering evenly from approximately 6 cm at the base to 1.6 cm at the tip (Figure 8). The tusk diameter increased evenly to the point of eruption at approximately 175 cm from the tip and then decreased rapidly within the skull. A regression of average outer diameter to distance from the tip for the length of tusk starting at the point of eruption FIGURE 7. (A) Three dimensional reconstruction made from CT data of the adult male dentition of M. monoceros showing the dorsal view of the fully developed tusk, the unerupted tusk, and the vestigial teeth, and (B) a lateral view of the complete head illustrating the location of the coronal slice at the level of the right vestigial tooth. 16_Nweela_pg223-240_Poles.indd 22916_Nweela_pg223-240_Poles.indd 229 11/18/08 9:13:28 AM11/18/08 9:13:28 AM 230 SMITHSONIAN AT THE POLES / NWEEIA ET AL. resulted in an equation yielding the average diameter in centimeters as follows: Diameter H11005 1.75 H11001 0.025 H11003 (dis- tance from tip), with an R 2 value of 0.989. The average pulp chamber diameter is shown in lower plot of the graph in Figure 8 and does not mirror the linear trend of the outer diameter. The chamber tapers evenly for the fi rst 100 cm from the tip and then reaches a plateau of approximately 1.75 cm for the next 50 cm and decreases in diameter slightly at 150 to 175 cm from the tip. The pulp diameter increases dramatically over the last 10 cm at base of the tusk. Soft tissue remnants were visible in the pulp chamber of all the frozen sections. A photo of an intact tusk with the entire body of pulp tissue removed is shown in the photograph in Figure 8. The cross sections of the tusk were often not sym- metric, but rather slightly oblong, with a major diameter in many sections being several millimeters larger than the minor diameter. The shape of the pulp chamber generally FIGURE 8. (top) A plot of the average cross sectional diameter in cm of the outer surface and pulp chambers of the sectioned tusk as a function of distance from the tip. (bottom) Photograph shows pulpal tissue removed from an intact tusk. 16_Nweela_pg223-240_Poles.indd 23016_Nweela_pg223-240_Poles.indd 230 11/18/08 9:13:47 AM11/18/08 9:13:47 AM CONSIDERATIONS OF NARWHAL DENTITION 231 mimicked the shape and profi le of the outer surface. The walls of the inner pulp chamber were very smooth when observed after removing the soft tissue remnants. The outer surface of the tusk demonstrated a series of major and minor ridges and valleys that progressed down the length of the tusk following a left-hand helix. The major ridges were anywhere from 2 to 10 mm in width and 1 to 2 mm in depth, while the minor ridges were ap- proximately 0.1 mm in width and depth. Brown and green deposits covered the major valleys, while the tops of the highest and broadest ridges were clean and white, accentu- ating the helical pattern of these features (Figure 9). The tip section was relatively smooth with an oblique, slightly concave facet approximately 3 cm in length ex- tending backward from the rounded end (Figure 10). A small stained deposit was present in the deepest depression of the facet. Higher magnifi cation did not reveal surface scratches indicating abrasive wear, and a small occluded remnant of the pulp chamber could be seen at the tip. The surface of the facet had what appeared to be many small grooves and smooth indentations on the surface that may be due to the exposure of softer areas in the layered tissue described later in the microanalysis. The lack of abrasion scratches, the concave profi le, and these small depressions indicate that the facet is more likely due to a combination of abrasion and erosion from abrasive slurry, such as sand, rather than being caused by rubbing against a hard abra- sive surface. Facets were found on many but not all tusks observed during the hunting season, while all exhibit the smoothly polished or clean tip section of approximately 10 cm in length. It was not possible to determine the ana- tomical orientation of the facet in the sectioned tusk, but fi eld observations describe it as varying in orientation from right, left, and downward, with an upward orienta- tion rarely being reported. Inuit descriptions of gross tusk morphology are de- scribed by variations of form within each sex and dimor- phic traits that differentiate tusk expression in females. Most hunters note that the blood and nerve supply in the pulp extends to the tip, and indeed, some hunters are ex- perienced in the extirpation of the pulp to its entire length. They describe a receding pulpal chamber for older nar- whal, which is a consistent fi nding with the increased age of most mammalian teeth. However, the female tusk is quite different in morphology. Female tusks are shorter, narrower, and evenly spiraled and denser, with little or no pulp chamber, even at an early age. Such an observation suggests a difference in functional adaptation as females FIGURE 9. A photograph showing the helical staining due to surface deposits of algae that remain in the deeper grooves of the tusk. 16_Nweela_pg223-240_Poles.indd 23116_Nweela_pg223-240_Poles.indd 231 11/18/08 9:14:00 AM11/18/08 9:14:00 AM 232 SMITHSONIAN AT THE POLES / NWEEIA ET AL. would lack substantial tusk innervation to sense their en- vironment. TUSK MICROMORPHOLOGY A mid-length section was chosen for visible and scan- ning electron microscopy (SEM). A 1-mm-thick slice was cut from the end of a section and polished using a metal- lographic polishing wheel and a series of abrasives down to 4000 grit size. Care was taken to retain the hydration of the section during processing and observation. The cross- sectional visual observation of this slice showed the tusk to be composed of several distinct layers (Figure 11). An outermost layer described as cementum was more trans- lucent and approximately 1.5 to 2 mm thick. There was a distinct boarder between this outermost layer and the underlying dentin. The bulk of the section was made up of a relatively homogenous dentin that had numerous dis- tinct rings, appearing much like the growth rings in a tree. The outermost rings were slightly whiter in color while the innermost ring adjacent to the pulp chamber appeared slightly darker opaque than the surrounding dentin rings. The dark lines radiating outward from the pulp chamber to the surface were associated with microtubules observ- able under SEM and are described later. An additional pie-shaped section was cut from this polished slice to include both the pulpal and outer sur- FIGURE 10. A close-up photograph of the fl at facet on the tip of the tusk. A translucent remnant of the pulp chamber can be seen at the tip as well as many minor grooves and indentations. 16_Nweela_pg223-240_Poles.indd 23216_Nweela_pg223-240_Poles.indd 232 11/18/08 9:14:03 AM11/18/08 9:14:03 AM CONSIDERATIONS OF NARWHAL DENTITION 233 faces for SEM observation. The specimen was cleaned to remove surface debris using a mild HCL acid etch fol- lowed by rinsing in dilute sodium hypochlorite. This piece was then dried in a vacuum dessicator and gold sputter coated for conductivity. The SEM images of the outer sur- face showed that the dark stain material found at the base of the grooves was made up of microscopic diatoms from algae deposits adhering in multiple layers to the surface (Figure 12). The smooth white ridge areas were regions where these deposits were either very sparse or completely miss- ing. The level of artifacts and debris on the surface made it diffi cult to observe the underlying tooth surface as cleaning did not remove the tightly adhered deposits in the grooves but did occasionally expose the ends of small tubule-like canals opening to the tusk surface. A more thorough cleaning removed more of the deposits and exposed the outer openings with regular frequency (Figure 13). These openings were approximately one to two micrometers in diameter and were similar in appearance to those found on the pulpal surface of the dentin. Scanning electron microscopy observation of the pulpal surface of the section revealed the openings of multiple den- tin tubules. These dentin tubules ranged from 0.5 to several micrometers in diameter and were evenly distributed with spacing of approximately 10 to 20 micrometers between tubules (Figure 14). This spacing was less dense than that observed on the pulpal surfaces of human or bovine teeth, FIGURE 11. A transilluminated corner of a polished cross section showing the many distinct layer or rings within the tissue. The wide band of tissue making up the outer surface is described as ?cementum.? 16_Nweela_pg223-240_Poles.indd 23316_Nweela_pg223-240_Poles.indd 233 11/18/08 9:14:05 AM11/18/08 9:14:05 AM 234 SMITHSONIAN AT THE POLES / NWEEIA ET AL. where spacing is approximately 3 to 5 micrometers between tubules. The appearance of the bell-shaped openings and lu- men of the tubules was similar to that observed in the teeth of other mammals. The cross-sectional surface of one pie- shaped piece was acid etched to remove the collagen smear layer that forms as an artifact of polishing. This section and other serial sections taken across the tubules revealed that the tubules radiated outward from the pulpal surface through the entire thickness of the dentin and appeared to communicate into the outermost cementum surface layer (Figure 15). This observation is in contrast to what is found in masticating teeth of mammals, where tubules radiate through the body of the dentin but terminate within dentin or at the base of the outer enamel layer. The fl exural strength of the dentin from two fresh sections, one close to the base and one mid length down the tusk, was measured using 2 H11003 2 H11003 15 mm rectangu- lar bars cut longitudinally down the length of the tusk. These bars were loaded to fracture in a three-point bend- ing mode over a 10-mm span using a universal testing ma- chine. Nine specimens were cut from the midsection of the tusk and four from the base section. The transverse rupture strength at the midsection was 94.6 H11006 7.0 MPa (mean H11006 standard deviation) and 165.0 H11006 11.7 MPa near the base. The bars from near the base underwent much more deformation prior to fracture than those from the midsection with approximately twice the amount of strain occurring at fracture. FIGURE 12. A scanning electron micrograph of the outside tusk surface showing stain deposits composed of diatoms and algae. 16_Nweela_pg223-240_Poles.indd 23416_Nweela_pg223-240_Poles.indd 234 11/18/08 9:14:07 AM11/18/08 9:14:07 AM CONSIDERATIONS OF NARWHAL DENTITION 235 DISCUSSION Imaging and dissection of adult male, adult female, and fetal narwhal specimens recorded a detailed visual re- cord of the cranial and dental anatomy. Among the fi nd- ings were three new discoveries of the dental anatomy and one observation of growth and development for the tusks. The fi rst major fi nding was the presence of paired vestigial teeth in all three specimens. Although a previous report in the literature found single vestigial teeth in a small collec- tion of narwhal skulls (Fraser, 1938), this is the fi rst study to document paired vestigial teeth. The lack of prior docu- mentation on vestigial teeth may be due to their location, as they were embedded in bone in the female specimen and suspended in the tissue located lateral to the anterior third of the maxillary bone plate. Radiography and digital im- aging provided an undisturbed view of these teeth in situ. The second discovery was linked to the anatomical loca- tion of all four maxillary teeth and their relative locations during growth and development as the two pairs of teeth reverse positions. In the fetus, the future tusks are located anteromedially to the vestigial teeth pair of teeth at four to FIGURE 13. A scanning electron micrograph at 1000X magnifi cation of the outer surface of the tusk after cleaning deposits from the surface. Tubule openings can be observed on the surface at a regular frequency. The large center orifi ce is approximately two micrometers in diameter. 16_Nweela_pg223-240_Poles.indd 23516_Nweela_pg223-240_Poles.indd 235 11/18/08 9:14:20 AM11/18/08 9:14:20 AM 236 SMITHSONIAN AT THE POLES / NWEEIA ET AL. six months in development. The fully developed tusks are located posteromedially to the vestigial pair of teeth in the adult narwhal. The third fi nding was evidence of devel- oped sockets for the vestigial teeth that extend posteriorly to the base of the developed tusks and communicate with their nerve and blood supply. Evidence of these developed vestigial tooth sockets suggests that this species may have exhibited at least two pairs of well-developed teeth at some point in its evolution. Likewise, if the vestigial teeth never existed beyond their current state, then well-developed sockets for these structures would not be expected as visu- alized in the fetal and adult female specimen. Intracranial dissection revealed fi fth cranial nerve pathways that were consistent with other mammals, though there were some expected modifi cations based on the skull asymmetry. The gross morphology of the male narwhal tusk showed a surprisingly unique feature by having a nearly full length pulp chamber. This observation is confi rmed by most of the Inuit interviewed, though this feature has dimorphic characteristics, as traditional knowledge de- scribes females with little to no pulp chamber, even at younger ages. This is much different from other tusked mammals, where the pulp chamber is often only a small proportion of the tusk length. It would also seem counter- intuitive for a tooth evolving in a harsh and cold environ- ment to contain vital vascular and nervous tissue through- FIGURE 14. A scanning electron micrograph of the pulpal wall showing the opening of a dentin tubule. The tubule orifi ce is approximately 1 micrometer in diameter. The size and shape of the tubules is similar to those found in human and other mammalian teeth. 16_Nweela_pg223-240_Poles.indd 23616_Nweela_pg223-240_Poles.indd 236 11/18/08 9:14:25 AM11/18/08 9:14:25 AM CONSIDERATIONS OF NARWHAL DENTITION 237 out its length. The pulp chamber also compromises the strength of this long and seemingly fragile tooth. The re- sidual chamber seen at the tip of the tusk indicates that the chamber forms throughout tusk growth and develop- ment. One conversation with a broker of harvested tusks revealed that occasionally, a tusk is seen where the pulp chamber is very small and narrow and this usually occurs in larger and older tusks. It was not possible to verify the order or timing of dentin deposition using the methods of this study, but the possibility exists that the inner pulpal layer of dentin may be a feature of dentin formed at a later stage of tooth development or as a process of aging. The left-hand helical nature of tooth development has been hypothesized to be a functional adaptation to main- tain the overall concentric center of mass during growth. This hypothesis certainly makes a great deal of sense when one considers the hydrodynamic loads that would develop if the tooth were curved or skewed to one side. The clean smooth tip and facet observed in this specimen appears to be a secondary feature resulting from some form of abra- sion and/or erosion. The lack of scratch patterns and the presence of large and small concavities across the facet in- dicate it is not formed by abrasion against a hard surface, but rather could result from gradual attrition by an abrasive slurry, such as sand or sediment. This cleanly polished tip appears on every tusk observed, regardless of the presence or absence of the facet. The behavior that causes this fea- ture must be almost universal and is likely to be continuous, FIGURE 15. A scanning electron micrograph of a polished section of dentin shows the radiating and continuous nature of the tubules. Serial cross sections of tubules confi rmed their presence throughout the tusk wall. 16_Nweela_pg223-240_Poles.indd 23716_Nweela_pg223-240_Poles.indd 237 11/18/08 9:14:42 AM11/18/08 9:14:42 AM 238 SMITHSONIAN AT THE POLES / NWEEIA ET AL. as the algae deposits that stain the surface would likely re- appear if not continually removed. Water turbulence alone would probably not account for removal of these deposits from the tip and ridge areas of the tusk. The cleaned ridges are also smoothed, indicating that they could be cleaned by physically rubbing against a surface such as ice. There have also been traditional knowledge descriptions of ?tusking,? where males will gather in small groups and rub tusks in a nonaggressive manner. Hunters clean harvested tusks by rubbing them with sand to remove these deposits. Perhaps tusks come in contact with sand and sediment when nar- whal feed close to the bottom. The cementum layer on the outer surface of the tusk is also a rare feature for an erupted tooth. Cementum is generally found as a transitional layer between dentin and the periodontal ligaments that hold teeth into bone. These ligaments are able to attach to the cementum with small fi bers, tying it to the surrounding bone. This appears to be consistent with the cementum observed in sections from the tusk base that were attached to segments of bone. In human teeth, however, if cementum becomes exposed to the oral environment, it is rapidly worn away, exposing the root dentin. In the tusk, the cementum layer appears to remain intact, even after long exposure to the ocean environment. The thickness of the cementum layer also appears to increase as the tooth increases in diameter. Cementum in mammalian teeth is generally a more pro- teinaceous tissue with greater toughness than enamel or dentin. A toughened outer layer that increases in thickness toward the tusk base would be consistent with the me- chanics of fracture resistance, where tensile stresses would also be highest at the tusk surface and base. This tough- ened layer of tissue would resist cracking under functional stresses that could lead to tusk fracture. It is impossible to tell from these studies what causes color changes that distinguish the rings observed within the dentin, but one possibility would be a change in devel- opmental growth conditions, such as nutrition (Nweeia et al., in press) (Figure 16). The fl exural strength and work of fracture both increased for dentin when comparing the tusk base to the midsection. The fl exural strengths of 95 MPa at mid tusk and 165 MPa at the base compare to approxi- mately 100 MPa for human dentin. These are adaptations for a tooth that must withstand high fl exural stresses and deformation rather than the compressive loads of chewing. The microanatomy of the tusk also provides insight into potential function. Scanning electron micrographs of the pulpal surface revealed tubule features that are similar in size and shape to the dentin tubules found in masticating teeth. The tubule diameters are similar to those observed in human teeth, but the spacing of these tubules across the pulpal wall is three to fi ve times wider than that seen in human dentin. The polished cross sections show that these tubules run continuously throughout the entire thickness of dentin, just as they do in human dentin. A surprising fi nding, however, was the presence of tubule orifi ces on the outer surface of the cementum. This indicates that the den- tin tubules communicate entirely through the wall of the tusk with the ocean environment. It is well established that dentin tubules in human and animal teeth provide sensory capabilities. Exposure of these tubules to the oral environ- ment in human teeth is responsible for sensing temperature changes, air movement, and the presence of chemicals, such as sugar. An example of this phenomenon would be the pain one senses in a cavity when the tooth is exposed to sugar or cold air. The decay from the cavity removes the overlying protective enamel and exposes the underly- ing dentin, allowing the dentin tubules to communicate directly with the oral cavity. Changes in temperature, air movement, or osmotic gradient set up by the sugar cause movement of fl uid within these tubules. This movement is detected by neurons at the pulpal end of the tubule, and these neurons send the pain signal to our brains. Narwhal teeth have similar physiology to human teeth, having both FIGURE 16. A section (2.0 cm corresponding to the third plotted point in Figure 8) cut near the tusk tip showing multiple dentin rings under transillumination. 16_Nweela_pg223-240_Poles.indd 23816_Nweela_pg223-240_Poles.indd 238 11/18/08 9:14:54 AM11/18/08 9:14:54 AM CONSIDERATIONS OF NARWHAL DENTITION 239 pulpal neurons and dentin tubules. The most distinguish- ing difference is that the tubules in the narwhal tusk are not protected by an overlying layer of enamel. This raises the distinct possibility that the tusk could provide a vari- ety of sensory capabilities. Any stimulus that would result in movement of fl uid within these tubules could possibly elicit a response (Figure 17). These include ion gradients, such as water salinity, pressure gradients caused by dive depth or atmospheric pressure changes, air temperature and movement, or possibly other chemical stimuli specifi c to food sources or environment. Field experiments on three captive male narwhal completed during August 2007 in the Canadian High Arctic provided evidence that water salinity is one stimulus that can be sensed by this tusk. Introduction of a high salt ion solution (approximately 42 psu), immediately after freshwater exposure, within a fi xed gasket isolating a 35-cm portion of tusk surface, produced a marked movement of the head region and co- ordinated respiratory response. Two separate salt ion so- lution stimuli in two males and one stimulus in the third male were witnessed by twelve team members. Responses subsided immediately after freshwater was reintroduced to the tusk gasket. Though monitoring equipment was at- tached to the whale during experimentation, physiologic data recordings (EEG and ECG) were hindered by diffi cult fi eld conditions. ACKNOWLEDGMENTS Our sincere and kind thanks, qujanamik, to the Inuit High Arctic communities of Nunavut and northwestern Greenland and the 55 elders and hunters who gave their time and Inuit Qaujimajatuqangit, knowledge to add to the understanding for this extraordinary marine mammal and its unique tusk. We also wish to acknowledge the sup- port of the National Science Foundation; Astromed?s Grass Telefactor Division for their gracious support in loaning electroencephalographic equipment; Arctic Research Di- vision, Fisheries and Oceans, Canada; the National Geo- graphic Society, The Explorers Club, World Center for Exploration, the National Institutes of Health?s (NIH) Imaging Department, and the Johns Hopkins University?s Integrated Imaging Center, as well as individuals, includ- ing Leslee Parr, San Jose State University; Sam Ridegway, Judith St. Ledger, and Keith Yip, Sea World, San Diego; Joseph Meehan; Jim (Wolverine) Orr; Giuseppe Grande; Glenn Williams, Director of Wildlife, Nunavut Tunngavik, Inc.; Doug Morris and John Butman, NIH MRI Research Facility; Sandie Black, Head of Veterinary Services, Cal- gary Zoo Animal Care Centre; Rune Dietz, Department of Arctic Environment, National Environmental Research Institute; Mosesie Kenainak; Mogens Andersen, Assistant Curator, Vertebrate Department, Zoological Museum of Copenhagen; Michel Gosselin and Natalia Rybczyn- ski, Canadian Museum of Nature; Judith Chupasko and Mark Omura, Museum of Comparative Zoology, Har- vard University; Lisa Marie Leclerc, University of Quebec at Rimouski; Kevin Hand, Newsweek Corporation; Bruce Crumley, Hawk Enterprises; and William Fitzhugh, Daryl Domning, and Dee Allen, National Museum of Natural History, Smithsonian Institution. EXPANDED AUTHOR INFORMATION Martin T. Nweeia, Harvard University, School of Den- tal Medicine, 188 Longwood Ave, Boston, MA 02115, USA; also Smithsonian Institution, Division of Mammals, Department of Zoology, 100 Constitution Avenue, Wash- ington, DC 20056, USA. Cornelius Nutarak and David Angnatsiak, Community of Mittimatilik, Nunavut X0A FIGURE 17. Tusk of male M. monoceros as a hydrodynamic sen- sor. The anatomic features of the narwhal tusk provide the potential for sensing stimuli that would result in movement of interstitial fl uid within the dentin tubules. This fl uid movement stimulates neurons located at cellular odontoblastic layer found at the base of each tu- bule. 16_Nweela_pg223-240_Poles.indd 23916_Nweela_pg223-240_Poles.indd 239 11/18/08 9:14:56 AM11/18/08 9:14:56 AM 240 SMITHSONIAN AT THE POLES / NWEEIA ET AL. 0S0, Canada. Frederick C. Eichmiller, Delta Dental of Wis- consin, 2801 Hoover Road, Stevens Point, WI 54481, USA. Naomi Eidelman, Anthony A. Giuseppetti, and Janet Quinn, ADAF Paffenbarger Research Center, National Institute of Standards and Technology, 100 Bureau Drive, Gaithers- burg, MD 20899-8546, USA. James G. Mead and Charles Potter, Smithsonian Institution, Division of Mammals, De- partment of Zoology, 100 Constitution Avenue, Washing- ton, DC 20056, USA. Kaviqanguak K?issuk and Rasmus Avike, Community of Qaanaaq, Greenland 3980. Peter V. Hauschka, Children?s Hospital, Department of Orthopae- dic Surgery, Boston, MA 02115, USA. Ethan M. Tyler, Na- tional Institutes of Health, Clinical Center, 9000 Rockville Pike, Bethesda, MD 20892, USA. Jack R. Orr, Fisheries and Oceans, Canada, Arctic Research Division, 501 University Crescent, Winnipeg, MB, R3T 2N6, Canada. Pavia Nielsen, Community of Uummannaq, Greenland 3962. LITERATURE CITED Beddard, F. E. 1900. A Book of Whales. New York: Putnam and Sons. Best, R. C. 1981. The Tusk of the Narwhal (L.): Interpretation of Its Function (Mammalia: Cetacea). Canadian Journal of Zoology, 59: 2386? 2393. Brear, K., J. D. Curry, C. M. Pond, and M. A. Ramsay. 1990. The Me- chanical Properties of the Dentine and Cement of the Tusk of the Narwhal Monodon monoceros Compared with Those of Other Mineralized Tissues. Archives of Oral Biology, 35(8):615? 621. Brown, R. 1868. Cetaceans of the Greenland Seas. Proceedings of the Zoological Society of London, 35: 552? 554. Bruemmer, F. 1993. The Narwhal Unicorn of the Sea. Emeryville: Pub- lishers Group West. Busch, F. 1890. Zur physiologie und pathologie der zahne des elefanten. Verhandlungen der Deutschen Odontologischen, Gesellschaft, 1: 246? 315. Clark, J. W. 1871. On the Skeleton of a Narwhal (Monodon monoceros) with Two Fully Developed Tusks. Proceedings of the Zoological Society of London, VI(2): 41? 53. Colyer, J. F. 1915. Injuries of the Jaws and Teeth in Animals. The Dental Record, 35: 61? 92, 157? 168. de Muizon, C., and D. P. Domning. 2002. The Anatomy of Obobenoce- tops (Delphinoidea, Mammalia), the Walrus-Like Dolphin from the Pliocene of Peru and Its Palaeobiological Implications. Zoological Journal of the Linnaean Society, 134: 423? 452. de Muizon, C., D. P. Domning, and M. Parrish. 1999. Dimorphic Tusks and Adaptive Strategies in a New Species of Walrus-Like Dolphin (Odobenocetopsidae) from the Pliocene of Peru. Earth and Plan- etary Sciences, 329: 449? 455. Eales, N. B. 1950. The Skull of the Foetal Narwhal, Monodon monoc- eros L. Philosophical Transactions of the Royal Society of London, Series B, Biological Sciences, 235(621): 1? 33. Ellis, R. 1980. The Book of Whales. New York: Alfred A. Knopf. Fraser, F. C. 1938. Vestigial Teeth in the Narwhal. Proceedings of the Linnaean Society of London, 150: 155? 162. Freuchen, P. 1935. Mammals. Part II. Field Notes and Biological Obser- vations. Report of the Fifth Thule Expedition, 1921? 24, 2(4? 5): 68? 278. Gervais, P. 1873. Remarques sur la Dentition du Narval. Journal de Zo- ologie, 2: 498? 500. Geist, O. W., J. W. Manley, and R. H. Manville. 1960. Alaskan Records of the Narwhal. Journal of Mammalogy, 41(2): 250? 253. Goethe, J. W. 1949. Gedenkausgabe der Werke, Brief und Gesprache. Volume 11. Z?rich: Artemis-Verlag. Harrison, R. J., and J. E. King. 1965. ?Family Monodontidae.? In Ma- rine Mammals, ed. R. J. Harrison and J. E. King, pp. 36? 39. Lon- don: Hutchison University Library. Hartwig, G. 1874. The Polar and Tropical Worlds. Ottawa: J. W. Lyon. Hay, K. A., and A. W. Mansfi eld. 1989. ?Narwhal? Monodon monoc- eros Linnaeus, 1758.? In Handbook of Marine Mammals, ed. S. H. Ridgeway and R. Harrison, Volume 4, pp. 145? 159. London: Aca- demic Press. Heyning, J. E. 1984. Functional Morphology Involved in Interspecifi c Fighting of the Beaked Whale, Mesoplodon carlhubbsi. Canadian Journal of Zoology, 645:59? 60. Kingsley, M., and M. A. Ramsay. 1988. The Spiral in the Tusk of the Narwhal. Arctic, 41(3): 236? 238. Lowe, A. P. 1906. The Cruise of the Neptune. Report on the Domin- ion Government Expedition to Hudson Bay and the Arctic Islands, 1903? 1904. Ottawa: Government Printing Bureau. Mansfi eld, A. W., T. G. Smith, and B. Beck.1975. The Narwhal (Mon- odon monoceros) in Eastern Canadian Waters. Journal of the Fish- eries Research Board of Canada, 32(7): 1041? 1046. Mead, J. G. 1989. ?Beaked Whales of the Genus Mesoplodon.? In Hand- book of Marine Mammals. Volume 4. River Dolphins and Larger Toothed Whales, ed. S. H. Ridgeway and R. Harrison, pp. 349? 430. London: Academic Press. Newman, M. A. 1971. Capturing Narwhals for the Vancouver Public Aquarium, 1970. Polar Record, 15: 922? 923. Nweeia, M. T., J. F. Thackeray, F. C. Eichmiller, P. Richard, L.-M. Leclerc, J. Lanham, and I. Newton. In press. A Note on Isotopic Analysis of Sectioned Tusks of Narwhal (Monodon monoceros) and Tusk Growth Rates. Annals of the Transvaal Museum. Nweeia, M. T., N. Eidelman, F. C. Eichmiller, A. A. Giuseppetti, Y. G. Jung, and Y. Zhang. 2005. Hydrodynamic Sensor Capabilities and Structural Resilience of the Male Narwhal Tusk. Abstract presented at the 16th Biennial Conference on the Biology of Marine Mam- mals, San Diego, Calif., 13 December 2005. Pederson, A. 1931. Mammals and Birds along the East Coast of Green- land. Fisheries Research Board of Canada Translation Series, 1206: 412? 417. Reeves, R. R., and E. Mitchell. 1981. The Whale behind the Tusk. Natu- ral History, 90(8): 50? 57. Roberge, M. M., and J. B. Dunn. 1990. Assessment of the Subsistence Harvest and Biology of Narwhal (Monodon monoceros L.) from Admiralty Inlet, Baffi n Island, Northwest Territories (Canada) 1983 and 1986? 89. Canadian Technical Report of Fisheries and Aquatic Sciences, 1947: 1? 32. Scoresby, W., Jr. 1820. An Account of the Arctic Regions, with a His- tory and Description of the Northern Whale-Fishery. Volumes 1? 2. London: Archibald Constable and Co. Silverman, H. B., and M. J. Dunbar. 1980. Aggressive Tusk Use by the Narwhal (Monodon monoceros L.). Nature, 284: 57. Thompson, D. W. 1952. On Growth and Form. Volume 2. 2nd ed. Lon- don: Cambridge University Press. Tomlin, A. G. 1967. Mammals of the U.S.S.R. Volume 9. Cetacea. Jeru- salem: Israel Program for Scientifi c Translation. Vibe, C. 1950. The Marine Mammals and Marine Fauna in the Thule District (N.W. Greenland) with Observations on Ice Conditions in 1939, 1940, and 1941. Meddelelser om Gr?nland, 150(6): 117. 16_Nweela_pg223-240_Poles.indd 24016_Nweela_pg223-240_Poles.indd 240 11/18/08 9:15:02 AM11/18/08 9:15:02 AM ABSTRACT. Approximately four decades ago, scientists were fi rst able to enter the un- dersea polar environment to make biological observations for a nominal period of time. The conduct of underwater research in extreme environments requires special consider- ation of diving physiology, equipment design, diver training, and operational procedures, all of which enable this under-ice approach. Since those fi rst ice dives in wetsuits and double-hose regulators without buoyancy compensators or submersible pressure gauges, novel ice diving techniques have expanded the working envelope based on scientifi c need to include the use of dive computers, oxygen-enriched air, rebreather units, blue- water diving, and drysuit systems. The 2007 International Polar Diving Workshop in Svalbard promulgated consensus polar diving recommendations through the combined international, interdisciplinary expertise of participating polar diving scientists, equip- ment manufacturers, physiologists and decompression experts, and diving safety offi cers. The National Science Foundation U.S. Antarctic Program scientifi c diving exposures, in support of underwater research, enjoy a remarkable safety record and high scientifi c productivity due to a signifi cant allocation of logistical support and resources to ensure personnel safety. INTRODUCTION Milestones of U.S. Antarctic diving activities (Table 1) start with the fi rst dive by Americans in Antarctic waters made just after New Year?s Day in 1947 as part of Operation Highjump, the United States? fi rst major postwar Ant- arctic venture. Lieutenant Commander Tommy Thompson and a Chief Dixon used ?Jack Brown? masks and Desco? oxygen rebreathers. Early scuba divers braved McMurdo Sound?s H110021.8?C water with wetsuits and double-hose regu- lators. Equipment advances since then have led to the use of variable volume drysuits, buoyancy compensators (BCs), and dive computers. Because of their resistance to freezing, however, double-hose regulators were used almost ex- clusively in the McMurdo area from 1963 until 1990. Since then, single-hose regulators that also resist freeze-up failure have been used. From 1947 to 1967, research diving operations fell under the control of the U.S. Naval Support Force Antarctica and divers adhered to established U.S. Navy diving regula- tions. In 1967, James R. Stewart, Scripps Institution of Oceanography diving offi cer, established guidelines for the conduct of research diving in the polar Michael A. Lang, Smithsonian Institution, Of- fi ce of the Under Secretary for Science, P.O. Box 37102, MRC 009, Washington, DC 20013-7012, USA. Rob Robbins, Raytheon Polar Services Company, 7400 South Tucson Way, Centennial, CO 80112, USA. Corresponding author: M. Lang (langm@si.edu). Accepted 28 May 2008. Scientifi c Diving Under Ice: A 40-Year Bipolar Research Tool Michael A. Lang and Rob Robbins 17_Lang_pg241-252_Poles.indd 24117_Lang_pg241-252_Poles.indd 241 11/17/08 9:54:07 AM11/17/08 9:54:07 AM 242 SMITHSONIAN AT THE POLES / LANG AND ROBBINS TABLE 1. Milestones of USAP Dive Program (adapted from Brueggeman, 2003). Date(s) Milestone 1947 First dive by Americans in Antarctic waters, LCDR Thompson and Chief Dixon, as part of Operation Highjump, using Jack Brown masks and Desco oxygen rebreathers 1951 First Antarctic open-circuit scuba dive 1947? 1967 Research diving operations under USN Support Force, Antarctica 1961? 1962 Verne E. Peckham (Donald E. Wohlschlag project, Stanford University) logged 35 science dives tended topside on occasion by Arthur Devries and Gerry Kooyman 1962? 1963 John S. Bunt (Donald E. Wohlschlag project, Stanford University) logged 7 science dives 1963? 1964 G. Carleton Ray (New York Zoological Society), Elmer T. Feltz and David O. Lavallee logged 10 scuba dives 1963? 1964 Gerald Kooyman started diving under ice with Weddell Seals with Paul K. Dayton tending topside 1963? 1964 Willard I. Simmonds (Jacques S. Zaneveld project, Old Dominion University) logged 45 tethered science dives 1964? 1965 Gerry Kooyman, Jack K. Fletcher and James M. Curtis logged 71 science dives 1965? 1966 David M. Bresnahan (NSF OPP) and Leonard L. Nero dived on Zaneveld?s project 1965? 1966 G. Carleton Ray, Michael A. deCamp, and David O. Lavallee diving with Weddell seals 1967 NSF-SIO agreement for polar research diving (James R. Stewart) 1968 Paul K. Dayton benthic ecology project divers Charles Gault, Gerry Kooyman, Gordon Robilliard. Dayton has logged over 500 hundred dives under McMurdo ice 1978? 1979 Dry Valley Lake diving: George F. Simmons, Bruce C. Parker and Dale T. Andersen 1987 USAP Guidelines for Conduct of Research Diving 1990 Double-hose regulators phased out in favor of single-hose regulators. 1992 AAUS Polar Diving Workshop (Lang, M.A and J.R. Stewart, eds.) 1995 RPSC on-site Scientifi c Diving Coordinator (Rob Robbins) 2001 NSF-Smithsonian Interagency Agreement for polar research diving (Michael A. Lang) 2003? 2007 Svalbard ice diving courses (Michael A. Lang) 2007 International Polar Diving Workshop, Svalbard (M.A. Lang and M.D.J. Sayer, eds.) 2008 Smithsonian/NSF ice-diving regulator evaluation project, McMurdo (Michael A. Lang, P. I.) regions for the National Science Foundation (NSF) Of- fi ce of Polar Programs (OPP). Since 1995, Rob Robbins, Raytheon Polar Services Company, has served as onsite scientifi c diving coordinator. In 2001, Michael A. Lang, director of the Smithsonian Scientifi c Diving Program, enacted an Interagency Agreement between the Smithso- nian Institution and the NSF for the management of the U.S. Antarctic Program (USAP) scientifi c diving program. As NSF OPP Diving Safety Offi cer (DSO), these respon- sibilities include, with the USAP Diving Control Board, promulgation of diving safety standards and procedures, evaluation and training of prospective divers, and authori- zation of dive plans. The USAP Standards for the Conduct of Scientifi c Diving (USAP, 1991) references the scientifi c diving standards published by the American Academy of Underwater Sciences (AAUS). Approximately half of the Principal Investigators (Table 2) are employees of AAUS organizational member institutions. The USAP research- ers understand that polar diving demands the acceptance of responsibility for an increased level of risk and diver preparation. Polar conditions are more rigorous and de- manding of scientifi c divers and their equipment than most other diving environments. Approximately 36 scientists dive each year through USAP and have logged more than 11,400 scientifi c ice dives since 1989 (Figure 1). Average dive times are 45 minutes; generally, no more than two dives are made per day within the no-decompression limits. The USAP scientifi c diving authorization process requires submission of information on diver training and history, depth certifi cation, diving fi rst aid training (Lang et al., 2007) and drysuit experi- ence. Minimum qualifi cation criteria for NSF diving au- thorization include: (a) a one-year diving certifi cation; (b) 50 logged open-water dives; (c) 15 logged drysuit dives; and, (d) 10 logged drysuit dives in the past six months. Somers (1988) described ice diver training curricula con- siderations. A pre-dive orientation and checkout dive(s) are done on site to ensure that the diver exhibits a satisfac- tory level of comfort under the ice with their equipment. Divers new to the Antarctic program are usually accompa- nying experienced Antarctic research teams and are thus mentored in an ?apprentice? mode. However, divers must 17_Lang_pg241-252_Poles.indd 24217_Lang_pg241-252_Poles.indd 242 11/17/08 9:54:08 AM11/17/08 9:54:08 AM SCIENTIFIC DIVING UNDER ICE 243 become profi cient with the gear and techniques they will be using prior to deployment. THE POLAR DIVING ENVIRONMENT ICE FORMATION Ice crystallization begins at the air-sea interface where the temperature differential is greatest. Because the air may be as much as 50?C colder than the water, heat conduction to the air from the water promotes the formation of ice. Under calm conditions, this congelation ice is composed of needles, small disks, and dendritic stars and will form a smooth sheet over the sea. When the freezing sea is sub- jected to wind and wave action, frazil ice crystals clump together into pancake ice (0.5 m to 2 m in diameter) that consists of roughly circular, porous slabs with upturned edges. If the water between them freezes, the ?pancakes? may solidify and join together. Otherwise, pancake ice continually interacts with wind, waves, and other ice to create complex, many-layered fl oes of pack ice. When the ice sheet, whether congelation or frazil ice in origin, be- comes a solid surface joined to the shoreline, it forms fast ice. Once the ice sheet is established, it continues to grow from beneath. Low-density seawater emanating from be- neath ice shelves and fl oating glaciers undergoes adiabatic supercooling. Platelet ice crystals form in this supercooled water and fl oat upward, accumulating in an initially loose and porous layer at the bottom of the surface ice sheet. This unfrozen platelet layer (1 cm to several m thick) con- tinually solidifi es by freezing, increasing the thickness of the ice sheet. The platelet layer forms a substrate for the TABLE 2. Principal Investigators and Co-PIs of USAP diving projects (1989? 2006). Investigator Project Amsler, C. University of Alabama, Birmingham* Baker, W. Florida Institute of Technology/University of South Florida* Barry, J. Monterey Bay Aquarium Research Institute* Bosch, I. SUNY-Geneseo Bowser, S. NY Dept. of Health-Wadsworth Center Conlan, K. Canadian Museum of Natural History Davis, R. Texas A&M University* Dayton, P. Scripps Institution of Oceanography* DeVries, A. University of Illinois-Urbana Doran, P. University of Illinois-Chicago Dunton, K. University of Texas-Austin* Harbison, R. Woods Hole Oceanographic Institution* Kaiser, H. N/A Kennicutt, M. University of Texas-Austin* Kim, S. Moss Landing Marine Laboratories* Kooyman, G. Scripps Institution of Oceanography* Kvitek, R. California State University, Monterey Bay Lang, M. Smithsonian Institution* Lenihan, H. University of North Carolina* Madin, L. Woods Hole Oceanographic Institution* Manahan, D. University of Southern California* Marsh, A. University of Delaware McClintock, J. University of Alabama, Birmingham* McFeters, G. Montana State University Miller, M. Exploratorium, San Francisco Moran, A. Clemson University Oliver, J. Moss Landing Marine Laboratories* Pearse, J. University of California, Santa Cruz* Ponganis, P. Scripps Institution of Oceanography* Quetin, L. University of California, Santa Barbara* Torres, J. University of South Florida* Wharton, R. University of Nevada-Desert Research Institute Wu, N. Mo Yung Productions *AAUS organizational member institution. FIGURE 1. (top) USAP dive summary, 1989? 2006. (bottom) USAP authorized diver summary, 1989? 2006. 17_Lang_pg241-252_Poles.indd 24317_Lang_pg241-252_Poles.indd 243 11/17/08 9:54:08 AM11/17/08 9:54:08 AM 244 SMITHSONIAN AT THE POLES / LANG AND ROBBINS growth of microbial communities dominated by microal- gae fed upon by amphipods and ice fi sh. Ice may also crys- tallize on the benthos. This anchor ice generally forms at depths of 15 m or less, attaching to rocks and debris? and even to live invertebrates. If enough ice forms on these objects, they will fl oat up and may become incorporated into the ice sheet. FAST ICE Diving conditions are usually associated with solid fast-ice cover for most of the austral diving season at Mc- Murdo Station (annual average thickness 2 m, multiyear 4 m), limited freezing at Palmer Station (under 30 cm), periodically in the Svalbard fjords (average 1 m), and the perennially ice-covered Dry Valley lakes (greater than 6 m; Andersen, 2007). A solid fast-ice cover provides a calm, surge-free diving environment and offers a stable working platform with no surface wave action. Fast-ice strength and thickness varies with time of year and ambient tem- perature affecting diving operational support. The under- ice topography varies dramatically at dive site, time of year, microalgal activity, ocean current, age of ice, and other oceanographic and physical factors. When viewed from below, a fast-ice sheet may appear relatively homogenous as a hard, fl at surface but in places can be punctuated by cracks and openings that appear as bright lines in an oth- erwise dark roof. If platelet ice is present, the underside of the ice appears rough and uneven. Areas of multiyear ice and thick snow cover are darker. Where pressure ridges and tidal cracks are present, the under-ice topography has more relief. Large and small chunks of broken ice may jut down into the water column in profusion, creating an environment reminiscent of cave diving. Brine channels or ice stalactites form as seawater cools and freezes and salt is excluded. This salt forms a supercooled brine solution that sinks because of its increased density and freezes the seawater around it resulting in a thin, hollow tube of ice stretching down from the underside of the ice sheet. These brine channels can reach several m in length and may ap- pear singly or in clusters. PACK ICE Fast-ice diving differs from pack-ice diving (Quetin and Ross, 2009, this volume), where broken ice cover usually eliminates the need to cut access holes for diving because of easy access to the surface. The pack-ice environment tends to be more heterogeneous than that of fast ice. Ice may be present in all stages of development and the fl oes themselves may vary in size, age, structure, and integrity. Pack-ice divers will fi nd themselves under an ever-shifting and dynamic surface and wave action and currents must be considered. At sites where the pack ice is forced against the shore and is solid but unstable, an access hole will have to be opened near shore in shallow water. Tidal fl uctuations may alter the size of dive holes or vary the water depth under the holes. UNDERWATER VISIBILITY In August and September in the McMurdo region, un- derwater visibility may range up to a record 300 m. As solar radiation increases during the austral summer, an annual plankton bloom develops and quickly diminishes visibility to as little as 1 m by late December. Other water visibility factors infl uencing the polar regions include gla- cial melt and wind and temperature conditions. Visibility in the open waters of the Antarctic Peninsula may vary from 300 m to less than 3 m, depending on plankton den- sities and sea state. As glacial or sea ice melts, the resulting water may form a brackish water lens over the seawater. Visibility within these lenses is markedly reduced, even when the visibility in the water is still good otherwise. It may be possible to lose sight of the entry hole even when divers are near the surface. POLAR DIVING OPERATIONS DIVE ACCESS THROUGH ICE Tidal action, currents, and other forces produce open cracks and leads that divers may use to enter the water. Divers working from USAP research vessels often use the leads cut by the vessel for their access to the water (Quetin and Ross, 2009, this volume). A hydraulically operated mobile drill can be used to cut 1.3 m diameter holes in ice that is over 5 m in thickness. In addition to the pri- mary dive hole, at least one safety hole is required. Hole melters consisting of coiled copper tubing fi lled with hot circulating glycol or alcohol are used to open a clean, 1 m diameter hole in the thick ice cap that covers the freshwa- ter Dry Valley Lakes (Andersen, 2007), taking from sev- eral hours to several days. Chain saws can also be used to cut an access hole through ice that is 15 to 60 cm thick. Access holes are cut into square or triangular shapes and made large enough to accommodate two divers in the wa- ter simultaneously. Another method is to use Jiffy drills that bore pilot holes in ice 15 to 30 cm thick and then saws can be used to cut a large dive hole between them; 17_Lang_pg241-252_Poles.indd 24417_Lang_pg241-252_Poles.indd 244 11/17/08 9:54:12 AM11/17/08 9:54:12 AM SCIENTIFIC DIVING UNDER ICE 245 attaching ice anchors to the chunks of ice allows for easy removal once they are sawed free. For ice from 15 to 25 cm thick, ice saws and breaker bars (2 m lengths of steel pipe or solid bar with a sharpened tip) are used to cut and break away the ice to form a hole. Divers enter the wa- ter through pack ice from shore, from an infl atable boat launched from shore or a research vessel, or from large ice fl oes or a fast-ice edge. If dive holes are required in ice thicker than 5 m or in ice out of range of the mobile drill, explosives may be nec- essary. However, the use of explosives is generally discour- aged for environmental reasons and requires several hours of clearing ice from the hole before a dive can be made. Fast-ice diving requires one or more safety holes in addition to the primary dive hole. During times of the year when air temperatures are extremely cold, dive holes freeze over quickly. Positioning a heated hut or other portable shelter over a dive hole will delay the freezing process. So- lar powered electric muffi n fans are used to blow warm air from near the ceiling of the hut to the ice hole through a plastic tube. Down lines must mark all holes available for use on each dive because safety holes that are allowed to freeze at the surface are hard to distinguish from viable holes while diving under the ice. DOWN LINES AND TETHERS A down line is required on all untethered dives con- ducted from fast ice or any other stable overhead envi- ronment with limited surface access. Specifi c down line characteristics and components are described by Lang and Robbins (2007). A minimum of one supervisor/tender per dive is re- quired. Because they are a critical part of the diving opera- tion and the fi rst responders in case of accident, tenders receive training in diving fi rst aid (Lang et al., 2007), radio use and communication procedures, scuba gear assembly, tether management, and vehicle or boat operation. Dives conducted under fast ice where there is a current, reduced visibility or open blue water, or where the water is too shallow to maintain visual contact with the dive hole, require individual diver tethers that are securely attached at the surface. Use of the T- or L-shaped tether system is not ideal, making line-pull communication signals diffi cult and tether entanglement a possibility. Surface tender train- ing is necessary to maintain enough positive tension on the tether line to immediately recognize line-pull signals from the safety diver, without impeding the activity or motion of the scientists working under the ice. The safety diver?s func- tion is to keep tethers untangled, watch for large predators and communicate via line-pull signals to the surface and other working divers. Other hole-marking techniques to further protect against loss of the dive hole are snow removal (straight lines radiating outward from the dive hole that are very visible from under water) and benthic ropes which con- sist of 30 m lines laid out by divers when they fi rst reach the benthos, radiating outward like the spokes of a wheel from a spot directly beneath the dive hole and marked so that the direction to the dive hole is clearly discernible. POLAR DIVING EQUIPMENT Members of the dive team take particular care in the selection and maintenance of polar diving equipment (Lang and Stewart, 1992; Lang and Sayer, 2007). Antarc- tic waters are among the coldest a research diver can ex- pect to experience (H110021.8?C in McMurdo Sound). In these temperatures, not all diving equipment can be expected to operate properly and freeze-ups may be more frequent. Diving under total ice cover also imposes safety consid- erations that are refl ected in the choice of gear. We have developed specifi c care and maintenance procedures to en- sure the reliability of life support equipment. Divers are required to have two fully independent regulators attached to their air supply whenever they are diving under a ceiling. Modifi ed Sherwood Maximus regulators (SRB3600 models, Figure 2) have been used successfully through the installation of a heat retention plate and adjustment of the intermediate pressure to 125 FIGURE 2. Sherwood Maximus SRB3600 second stage with heat retention plate. 17_Lang_pg241-252_Poles.indd 24517_Lang_pg241-252_Poles.indd 245 11/17/08 9:54:13 AM11/17/08 9:54:13 AM 246 SMITHSONIAN AT THE POLES / LANG AND ROBBINS psi. These units are rebuilt at the beginning of each season and with more than 7,000 dives have a freeze-up incident rate of 0.3 percent. Proper use and pre- and post-dive care substantially improves the reliability of ice diving regu- lators, which must be kept warm and dry before a dive. Divers should not breathe through the regulator before submersion except to briefl y ensure that the regulator is functioning because of ice crystallization on the air deliv- ery mechanism from breath moisture. This is particularly important if the dive is being conducted outside in very cold air temperatures. During a dive, a regulator is never used to fi ll a lift bag (small ?pony bottles? are available for this purpose) because large volumes of air exhausted rapidly through a regulator will almost certainly result in a free-fl ow failure. Infl ator hoses are attached to the backup regulator in case the air supply to the primary regulator must be turned off to stem a free fl ow. The backup regu- lator second stage is attached to the cylinder harness or buoyancy compensator (BC) such that it is readily acces- sible and easily detached. If the second stage is allowed to hang loosely from the cylinder and drag on the bot- tom, it will become contaminated with mud and sediment and may not function properly when required. After the dive, the regulators are rinsed and allowed to dry. During rinsing, care is taken to exclude water from the interior regulator mechanism. The diver ensures that the regulator cap is seated tightly, that the hoses and plugs on the fi rst stage are secure, and that the purge on the second stage is not accidentally depressed during the rinse. The primary cause of regulator free-fl ow failure is from water entry within the mechanism that freezes once the regulator is used (Clarke and Rainone, 1995). Freshwater in the regu- lator may freeze simply with submersion of the regulator in seawater or upon exposure to extremely cold surface air temperatures. If multiple dives are planned, it is rec- ommended to postpone a freshwater rinse of the regulator until all dives are completed for the day. Infl ator valves are also subject to free-fl ow failure, be- cause of water entry into the infl ation mechanism. Drysuit and BC infl ators must be kept completely dry and hose connectors blown free of water and snow before attach- ment to the valve. When infl ating a drysuit or a BC, fre- quent short bursts of air are used. Infl ator buttons must never be depressed for longer than one second at a time because rapid air expansion, adiabatic cooling (5?C drop), and subsequent condensation and freezing may cause a free fl ow. Buoyancy compensators need to allow unimpeded ac- cess to drysuit infl ator and exhaust valves. Water must be removed from the BC bladder after diving and rinsing be- cause freshwater in the bladder may freeze upon submer- sion of the BC in ambient seawater. In the McMurdo area, BC use is not currently required when the dive is conducted under a fast-ice ceiling because of the lack of need for sur- face fl otation. A BC must never be used to compensate for excess hand-carried weight. Because of their buoyancy characteristics and durability in cold temperatures, steel, instead of aluminum, scuba cylinders are used. Divers must wear suffi cient weight, without over- weighting, to allow for maintenance of neutral buoyancy with a certain amount of air in the drysuit. Runaway nega- tive buoyancy is as great a safety problem to recover from as out-of-control ascent. Because of the amount of weight (30 to 40 lbs) and potential for accidental release, weight belts are not used. Diving Unlimited International (DUI) has developed weight and trim systems (Fig. 3) that retain the benefi ts of a harness while still allowing full or partial FIGURE 3. DUI weight and trim system with bilaterally remov- able weight pockets (by pulling surgical tubing loops) and shoul- der harness. 17_Lang_pg241-252_Poles.indd 24617_Lang_pg241-252_Poles.indd 246 11/17/08 9:54:15 AM11/17/08 9:54:15 AM SCIENTIFIC DIVING UNDER ICE 247 dumping of weight under water. The weight system pre- vents accidental release and improves comfort by shifting the weight load from the diver?s hips to the shoulders. Drysuit choice depends on the diver?s preference, the requirement for range and ease of motion, and the options available with each suit. Vulcanized rubber suits must be used when diving in contaminated water because of post- dive suit decontamination requirements. Drysuits must be equipped with hands-free, automatic exhaust valves. Over- infl ation of the drysuit should never be used as a means to compensate for excess hand-carried weight. The choice of drysuit underwear is perhaps more important than the choice of drysuit construction material, because it is the underwear that provides most of the thermal protection. Many divers wear an underlayer of expedition-weight poly- propylene with an outer layer of 400 g Thinsulate?. Dry gloves or mitts with an inner liner instead of wet gloves are now used with the drysuit. The DUI zipseal dry gloves enjoy widespread use and are effective at warm air equalization from the drysuit into the glove at depth. A disadvantage of these dry-glove systems is the complete lack of thermal pro- tection if the gloves fl ood or are punctured, and the related inevitability of fl ooding the entire drysuit. Severe cold can damage o-ring seals exposed to the environment requiring frequent cleaning and lubrication. Compressor care and adequate pre-operation warming are necessary to ensure a reliable supply of clean air checked by air-quality tests conducted at six-month intervals. Air fi lters and crankcase oil are scheduled to be changed on a regular basis. The fi ltering capacity of portable compressors is usually limited, necessitating air intake hose positioning upwind and well away from compressor engine exhaust. Manual condensate drains are purged frequently to prevent moisture contamination and freezing of the fi lter. Each diver conducts a functional check of all equip- ment before each dive. Particular attention is paid to regu- lators and infl ator valves. If leakage or free fl ow is detected at the surface, the dive is postponed and the gear serviced because it will certainly free fl ow at depth. All divers must be able to disconnect, with gloved hands, the low-pressure hose from a free-fl owing drysuit infl ator valve to avoid an uncontrolled ascent. Because a drysuit must be infl ated to prevent ?suit squeeze? with increasing pressure, it is most effi cient to regulate buoyancy at depth by the amount of air in the drysuit. Drysuits must be equipped with a hands-free ex- haust valve (Lang and Egstrom, 1990). The BCs are con- sidered emergency equipment, to be used only in the event of a drysuit failure. This procedure eliminates the need to vent two air sources during ascent, reduces the chance of BC infl ator free-fl ow, and simplifi es the maintenance of neutral buoyancy during the dive. The main purpose of air in a drysuit is to provide thermal insulation as a low- conductivity gas. Buoyancy compensators and drysuits must never be used as lift bags. When heavy items must be moved underwater, separate lift bags designed specifi cally for that purpose are used. Lang and Stewart (1992) con- cluded that there may be occasions when the drysuit diver is more at risk with a BC than without one. Accordingly, BCs are not required for dives under fast ice where a down line is deployed and the dive is not a blue-water dive. SURFACE-SUPPLIED DIVING UNDER ICE Robbins (2006) described USAP?s surface-supplied diving activities, history, equipment, training, operations, and costs. By taking advantage of the equipment and ex- pertise brought to the USAP program by commercial div- ers, scientifi c diving has benefi ted from the use of surface- supplied diving techniques. Safety, comfort, and effi ciency are enhanced in some applications by using this mode long associated with industry but rarely used in the scientifi c arena. Since 1992, USAP has supported surface- supplied diving. In that period, 459 surface-supplied dives (of 8,441 total dives) were logged by 32 divers (of 107 total divers). The vast majority of surface-supplied dives were performed by 8 divers. The USAP?s experience with EXO-26 masks has been 11 free-fl ows in 106 dives (10.4 percent failure rate). AGA masks have had 2 free-fl ows in 26 dives (7.7 percent fail- ure rate). These data come from dives in the Dry Valley Lakes where water temperatures range between 0?C and 2?C. The failure rate would be even higher in H110021.8?C wa- ter of McMurdo Sound. A minimum of two familiarization dives are made by each new surface-supplied diver over two days in addition to topside and underwater training. A three-person crew is the minimum personnel requirement including a supervi- sor/tender, a diver, and a suited standby diver using either scuba or surface supply. Currently, the majority of surface-supplied diving is done utilizing 2-m tall high-pressure gas cylinders as an air source. A large 35 cfm/150 psi diesel compressor and smaller 14 cfm/125 psi gas compressor are available but used rarely for scientifi c diving operations. The USAP uses Kirby-Morgan Heliox-18 bandmasks and Superlite-17 helmets. While these units have a greater propensity to freeze and free-fl ow than Sherwood Maximus scuba regu- lators, their track record is as good as either the EXO-26 or AGA Divator full-face masks. 17_Lang_pg241-252_Poles.indd 24717_Lang_pg241-252_Poles.indd 247 11/17/08 9:54:20 AM11/17/08 9:54:20 AM 248 SMITHSONIAN AT THE POLES / LANG AND ROBBINS POLAR DIVING HAZARDS AND EMERGENCIES FAST-ICE DIVING HAZARDS Lighting is often dim under a solid ice cover, particu- larly early in the austral spring when the sun is low on the horizon. The amount of snow cover and ice thickness will also attenuate light transmission. Microalgal blooms and increasing zooplankton during the austral summer reduces available light, making it diffi cult for divers to locate bud- dies, down lines, and underwater landmarks. High visibil- ity early in the austral summer season may make under-ice or benthic objects seem closer than they are. This illusion may entice divers to travel farther from the access hole than is prudent. The greatest hazard associated with fast-ice diving is the potential loss of the dive hole or lead. Access holes, leads, and cracks in the ice are often highly visible from below because of downwelling daylight streaming through them. However, dive holes may be diffi cult to see due to conditions of darkness or of covering the holes with por- table shelters. Therefore, a well-marked down line is re- quired for fast-ice dives. Divers maintain positive visual contact with the down line during the dive and avoid be- coming so distracted by their work that they fail to take frequent note of their position in relation to the access hole or lead. Problems requiring an emergency ascent are serious, since a vertical ascent is impossible except when a diver is directly under the dive hole or lead. Additional safety holes ameliorate the danger of losing the primary dive hole but former dive holes that have frozen over may still look like safety holes from below. To eliminate confu- sion in a frequently drilled area, all active holes are marked with a down line. PACK-ICE DIVING HAZARDS Pack ice is inherently unstable and its conditions can change rapidly, primarily from surface wind conditions. An offshore wind may blow pack ice away from the shore- line and loosen the pack, whereas an onshore wind may move signifi cant quantities of pack ice against shorelines or fast-ice edges, obstructing what may have been clear access areas when divers entered the water. Similarly, in- creased wind pressure on pack ice may make driving and maneuvering an infl atable Zodiac more diffi cult or impos- sible. Under a jumble of pack ice, the topography is remi- niscent of cave diving. The condition of the pack must be continually monitored by divers and tenders for changes that may affect dive safety and the entry area must be kept clear. Down lines and tethers can be disturbed by shifting pack ice, forcing dive tenders to be alert in keeping these lines free of moving ice. Surface swells, even if only light to moderate, may cause pack ice to oscillate up and down. In shallow water, it is possible for a diver to be crushed between rising and falling pack ice and the benthos. At Palmer Station, surges from the calving glacier in Arthur Harbor may create a similar hazard. Divers avoid diving under pack ice if the clearance between the ice and the benthos is 3 m or less. In addition, lighting may be dim under a heavy pack-ice cover. Open water develops in McMurdo Sound when the fast ice breaks up in late December or early January. In the Palmer region, any existing fast ice usually breaks up by the end of October. Pack ice may be present for another month or two, and intermittently after that, but open wa- ter generally characterizes the diving environment after early December. Kongsfjorden in Svalbard has not formed a substantial ice cover since 2005. Climatic conditions will cause variation in annual ice conditions. Divers operating in open water and from small boats fl y a ?diver down? or ?Alpha? fl ag to warn other boat traf- fi c in the area. When diving from small boats a rapid exit from the water into the boat may be necessary. Because this can be diffi cult when fully laden with gear, lines with clips hang over the side of the boat to temporarily secure gear and a ladder facilitates diver exit. When diving in blue water (a deep open water envi- ronment devoid of visual cues as to the diver?s vertical po- sition in the water column) blue water diving guidelines generally apply (Haddock and Heine, 2005). Divers are tethered and wear buoyancy compensators and a down line is deployed if conditions warrant. Divers operat- ing under pack ice in blue water often perceive current increases. Wind action causes the pack to move, which in turn moves the water directly below it. This effect de- creases with depth, such that divers in still water at 10 m will have the illusion of movement as the pack ice above them drifts. Ice-edge diving is usually conducted in blue water, and it tends to be shallow (less than 10 m). The underside of the ice sheet provides a depth reference lacking in ice-free blue water dives. Divers watch continuously for leopard seals known to lunge out of the water to attack people at the ice edge. They may also lurk under the ice waiting for a penguin, or a diver, to enter the water. If penguins in the area demonstrate a reluctance to enter the water, it may be an indication that a leopard seal is nearby. 17_Lang_pg241-252_Poles.indd 24817_Lang_pg241-252_Poles.indd 248 11/17/08 9:54:20 AM11/17/08 9:54:20 AM SCIENTIFIC DIVING UNDER ICE 249 MARINE LIFE HAZARDS Few polar animal species are considered dangerous to the diver. Southern elephant seals (Mirounga leonina) and Antarctic fur seals (Arctocephalus gazelli) may become aggressive during the late spring/early summer breeding season. Crabeater seals (Lobodon carcinophagus) have demonstrated curiosity toward divers and aggression to humans on the surface. Leopard seals (Hydrurga leptonyx) have been known to attack humans on the surface and have threatened divers in the water. A case report of the single known in-water fatality caused by a leopard seal is described by Muir et al. (2006). Should any aggressive seal approach divers in the water, similar techniques to those protecting against sharks are applied. Polar bears (Ursus maritimus) and walrus (Odobenus rosmarus) in the Arctic are considered predatory mammals against which diving personnel must be safeguarded. Encounters with all of the aforementioned mammals are usually restricted to areas of open water, ice edges, or pack ice. Divers in the fast ice around McMurdo may encoun- ter Weddell seals (Leptonychotes weddelli) in the water. Occasionally a Weddell seal returning from a dive may surface to breathe in a dive hole to replenish its oxygen stores after a hypoxic diving exposure (Kooyman, 2009, this volume). Usually the seal will vacate the hole once it has taken a few breaths particularly if divers are ap- proaching from below and preparing to surface. Divers must approach such a seal with caution, since an oxygen- hungry seal may aggressively protect its air supply. Weddell seals protecting their surface access will often invert into a head-down, tail-up posture to watch for ri- vals. Divers entering or exiting the water are particularly vulnerable to aggressive male Weddells, who tend to bite each other in the fl ipper and genital regions. There are no recorded incidents of killer whale (Orcinus orca) attacks on divers. POLAR DIVING EMERGENCIES The best method to mitigate scuba diving emergencies is through prevention. Divers must halt operations any time they become unduly stressed because of cold, fatigue, nervousness, or any other physiological reason. Similarly, diving is terminated if equipment diffi culties occur, such as free-fl owing regulators, tether-system entanglements, leaking drysuits, or buoyancy problems. Emergency situa- tions and accidents stem rarely from a single major cause and they generally result from the accumulation of several minor problems. Maintaining the ability to not panic and to think clearly is the best preparation for the unexpected. Most diving emergencies can be mitigated by assistance from the dive buddy, reinforcing the importance of main- taining contact between two comparably equipped scien- tifi c divers while in the water. Loss of contact with the dive hole may require div- ers to retrace their path. Scanning the water column for the down line is done slowly and deliberately because the strobe light fl ash rate is reduced in the cold water. If the hole cannot be found, an alternate access to the surface may have to be located. Often there will be open cracks at the point where fast ice touches a shoreline. Lost divers will have to constantly balance a desirable lower air consumption rate in shallow water with the need for the wider fi eld of vision available from deeper water. Maintaining a safe proximity to the surface ac- cess point has made losing the dive hole an extremely unlikely occurrence. Loss of the tether on a fast-ice dive that requires its use is one of the most serious polar diving emergencies. Lost diver search procedures are initiated immediately (i.e., assumption of a vertical position under the ice where the tethered buddy will swim a circular search pattern just under the ceiling to catch the untethered diver). The dan- ger associated with the loss of a tether in low visibility is mitigated if the divers have previously deployed a series of benthic lines. If a diver becomes disconnected from the tether down current under fast ice, it may be necessary to crawl along the bottom to the down line. To clearly mark the access hole divers deploy a well-marked down line, establish recognizable ?landmarks? (such as specifi c ice formations) under the hole at the outset of the dive, leave a strobe light, a fl ag, or other highly visible object on the substrate just below the hole or shovel surface snow off the ice in a radiating spoke pattern that points the way to the dive hole. The under-ice platelet layer can be several meters thick and can become a safety concern if positively buoy- ant divers become trapped within this layer, become dis- oriented, and experience diffi culty extricating themselves. The most obvious solution is to exhaust air from the dry- suit to achieve negative buoyancy. If this is not possible and the platelet layer is not too thick, the diver may stand upside down on the hard under surface of the ice so that the head is out of the platelet ice to orient to the posi- tion of the dive hole and buddy. Another concern is that abundant platelet ice dislodged by divers will fl oat up and plug a dive hole. 17_Lang_pg241-252_Poles.indd 24917_Lang_pg241-252_Poles.indd 249 11/17/08 9:54:21 AM11/17/08 9:54:21 AM 250 SMITHSONIAN AT THE POLES / LANG AND ROBBINS Fire is one of the greatest hazards to any scientifi c operation in polar environments. The low humidity ulti- mately renders any wooden structure susceptible to com- bustion and once a fi re has started, it spreads quickly. Dive teams must always exercise the utmost care when using heat or open fl ame in a dive hut. If divers recognize during the dive that the dive hut is burning they must terminate the dive and ascend to a safety hole or to the under surface of the ice next to the hole (but not below it) in order to conserve air. ENVIRONMENTAL PROTECTION There are research diving sites in Antarctica (e.g., Palmer sewage outfall, McMurdo sewage outfall, and Win- ter Quarters Bay) that must be treated as contaminated wa- ter environments because of the high levels of E. coli bac- teria (that have been measured up to 100,000/100 ml) or the presence of a hydrogen-sulfi de layer (e.g., Lake Vanda). Diving with standard scuba or bandmask, where a diver may be exposed to the water, is prohibited in these areas. Surface-supplied/contaminated-water diving equipment is used at these sites ranging from Heliox-18 bandmasks for use with a vulcanized rubber drysuit to Superlite-17 helmets that mate to special Viking suits. All researchers must avoid degrading the integrity of the environment in which they work. In particular, polar divers should avoid over-collecting, to not deplete an organism?s abundance and alter the ecology of a research site; unduly disturbing the benthos; mixing of water layers such as halo- clines; using explosives for opening dive holes; and, spilling oil, gasoline, or other chemicals used with machinery or in research. Increased attention to Antarctic Treaty protocols on environmental protection and implementation of the Antarctic Conservation Act have made human? seal inter- actions a more sensitive issue. Dive groups should avoid Weddell seal breeding areas during the breeding season and their breathing holes in particular. PHYSIOLOGICAL CONSIDERATIONS COLD Cold ambient temperature is the overriding limiting factor on dive operations, especially for the thermal protec- tion and dexterity of hands. Dives are terminated before a diver?s hands become too cold to effectively operate the dive gear or grasp a down line. This loss of dexterity can oc- cur quickly (5? 10 min if hands are inadequately protected). Grasping a camera, net, or other experimental apparatus will increase the rate at which a hand becomes cold. Switch- ing the object from hand to hand or attaching it to the down line may allow hands to rewarm. Dry glove systems have greatly improved thermal protection of the hands. The cold environment can also cause chilling of the diver, resulting in a reduced cognitive ability with pro- gressive cooling. Monitoring the progression of the fol- lowing symptoms to avoid life-threatening hypothermia is important: cold hands or feet, shivering, increased air consumption, fatigue, confusion, inability to think clearly or perform simple tasks, loss of memory, reduced strength, cessation of shivering while still cold, and fi nally hypo- thermia. Heat loss occurs through inadequate insulation, ex- posed areas (such as the head under an inadequate hood arrangement), and from breathing cold air. Scuba cylinder air is initially at ambient temperature and chills from ex- pansion as it passes through the regulator. Air consump- tion increases as the diver cools, resulting in additional cooling with increased ventilation. Signifi cant chilling also occurs during safety stops while the diver is not moving. Polar diving requires greater insulation, which results in decreasing general mobility and increasing the potential for buoyancy problems. This also means that an increased drag and swimming effort, along with the donning and doffi ng of equipment, all increase fatigue. SURFACE COLD EXPOSURE Dive teams are aware that the weather can change quickly in polar environments. While they are in the fi eld, all divers and tenders have in their possession suffi cient cold-weather clothing for protection in any circumstance. Possible circumstances include loss of vehicle power or loss of fi sh hut caused by fi re. Boat motor failure may strand dive teams away from the base station. Supervisors/ten- ders on dives conducted outside must also be prepared for the cooling effects of inactivity while waiting for the div- ers to surface. In addition, some food and water is a part of every dive team?s basic equipment. Besides serving as emergency rations, water is important for diver rehydra- tion after the dive. HYDRATION Besides the dehydrating effect of breathing fi ltered, dry, compressed air on a dive, Antarctica and the Arctic are extremely low-humidity environments where dehydra- tion can be rapid and insidious. Continuous effort is ad- vised to stay hydrated and maintain proper fl uid balance. 17_Lang_pg241-252_Poles.indd 25017_Lang_pg241-252_Poles.indd 250 11/17/08 9:54:21 AM11/17/08 9:54:21 AM SCIENTIFIC DIVING UNDER ICE 251 Urine should be copious and clear and diuretics (coffee, tea, and alcohol) should be avoided before a dive. DECOMPRESSION Mueller (2007) reviewed the effect of cold on de- compression stress. The relative contributions of tissue N 2 solubility and tissue perfusion to the etiology of de- compression sickness (DCS) are not resolved completely. Over-warming of divers, especially active warming of cold divers following a dive, may induce DCS. Divers in polar environments should, therefore, avoid getting cold during decompression and/or after the dive and if they feel hypo- thermic, wait before taking hot showers until they have rewarmed themselves, for example, by walking. The effect of cold on bubble grades (as measured by Doppler scores) may be the same for a diver who is only slightly cold as for one who is severely hypothermic. Long-term health ef- fects for divers with a high proportion of coldwater dives should be considered in the future. Dive computers were examined for use by scientifi c divers (Lang and Hamilton, 1989) and have now been ef- fectively used in scientifi c diving programs for almost two decades in lieu of U.S. Navy or other dive tables. Currently, the decompression status of all USAP divers is monitored through the use of dive computers ( UWATEC Aladin Pro) and data loggers (Sensus Pro). Battery changes may be needed more frequently because of higher discharge rates in extreme cold. Advantages of dive computers over tables include their display of ascent rates, no- decompression time remaining at depth, and their dive profi le down- loading function. Generally, no more than two repetitive dives are conducted to depths less than 130fsw (40msw) and reverse dive profi les for no- decompression dives less than 40msw (130fsw) and depth differentials less than 12 msw (40fsw) are authorized (Lang and Lehner, 2000). Oxygen-enriched air ( nitrox) capability (Lang, 2006) and rebreather use have, to date, not been requested nor implemented by the USAP Diving Program. Cold and the physical exertion required to deal with heavy gear in polar diving can increase the risk of DCS. Furthermore, because of the polar atmospheric effect, the mean annual pressure altitude at McMurdo Station is 200 meters. Un- der certain conditions, pressure altitude may be as low as 335 meters at sea level. Surfacing from a long, deep dive (on dive computer sea level settings) to an equivalent alti- tude of 335 meters may increase the probability of DCS. Safety stops of three to fi ve minutes between 10- to 30- foot (3.3 to 10 m) depths are required for all dives (Lang and Egstrom, 1990). ACKNOWLEDGMENTS The authors wish to thank the Smithsonian Offi ce of the Under Secretary for Science, the National Science Foundation Offi ce of Polar Programs, the U.K. NERC Fa- cility for Scientifi c Diving, Diving Unlimited International, Inc., and Raytheon Polar Services Company. LITERATURE CITED Andersen, D. T. 2007. ?Antarctic Inland Waters: Scientifi c Diving in the Perennially Ice-Covered Lakes of the McMurdo Dry Valleys and Bunger Hills.? In Proceedings of the International Polar Diving Workshop. ed. M. A. Lang and M. D. J. Sayer, pp. 163? 170. Wash- ington, D.C.: Smithsonian Institution. Brueggeman, P. 2003. Diving under Antarctic Ice: A History. Scripps Institution of Oceanography Technical Report. http:// repositories .cdlib .org/ sio/ techreport/ 22 (accessed 8 August 2008). Clarke, J. R., and M. Rainone. 1995. Evaluation of Sherwood Scuba Regulators for Use in Cold Water. U.S. Navy Experimental Diving Unit Technical Report 9? 95. Panama City: U.S. Navy. Haddock, S. H. D., and J. N. Heine. 2005. Scientifi c Blue-Water Diving. California Sea Grant College Program. La Jolla, Calif.: University of California. Kooyman, G. 2009. ?Milestones in the Study of Diving Physiology: Ant- arctic Emperor Penguins and Weddell Seals.? In Smithsonian at the Poles: Contributions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 265? 270. Washington, D.C.: Smithsonian Institution Scholarly Press. Lang, M. A. 2006. The State of Oxygen-Enriched Air (Nitrox). Journal of Diving and Hyperbaric Medicine, 36(2): 87? 93. Lang, M. A., and G. H. Egstrom, eds. 1990. Proceedings of the Biome- chanics of Safe Ascents Workshop. Costa Mesa, Calif.: American Academy of Underwater Sciences. Lang, M. A., and R. W. Hamilton, eds. 1989. Proceedings of the Dive Computer Workshop. Costa Mesa, Calif.: American Academy of Underwater Sciences and California Sea Grant College Program. Lang, M. A., and C. E. Lehner, eds. 2000. Proceedings of the Reverse Dive Profi les WorkshopWashington, D.C.: Smithsonian Institution Lang, M. A., A. G. Marsh, C. McDonald, E. Ochoa, and L. Penland. 2007. ?Diving First Aid Training for Scientists.? In Diving for Sci- ence: Proceedings of the AAUS 25th Symposium, ed. J. M. Godfrey and N. W. Pollock., pp. 85? 102. Dauphin Island, Ala.: American Academy of Underwater Sciences. Lang, M. A., and R. Robbins. 2007. ?USAP Scientifi c Diving Program.? In Proceedings of the International Polar Diving Workshop, ed. M. A. Lang and M. D. J. Sayer, pp. 133? 155. Washington, D.C.: Smith- sonian Institution. Lang, M. A., and M. D. J. Sayer, eds. 2007. Proceedings of the Inter- national Polar Diving Workshop. Washington, D.C.: Smithsonian Institution. Lang, M. A., and J. R. Stewart, eds. 1992. Proceedings of the Polar Div- ing Workshop. Costa Mesa, Calif.: American Academy of Under- water Sciences. Mueller, P. H. J. 2007. ?Cold Stress as Decompression Sickness Factor.? In Proceedings of the International Polar Diving Workshop, ed. M. A. Lang and M. D. J. Sayer, pp. 63? 72. Washington, D.C.: Smithsonian Institution. Muir, S. F., D. K. A. Barnes, and K. Reid. 2006. Interactions between Hu- mans and Leopard Seals. British Antarctic Survey Technical Report. Cambridge, U.K.: British Antarctic Survey. 17_Lang_pg241-252_Poles.indd 25117_Lang_pg241-252_Poles.indd 251 11/17/08 9:54:22 AM11/17/08 9:54:22 AM 252 SMITHSONIAN AT THE POLES / LANG AND ROBBINS Quetin, L. B., and R. M. Ross. 2009. ?Life under Antarctic Pack Ice: A Krill Perspective.? In Smithsonian at the Poles: Contributions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 285? 298. Washington, D.C.: Smithsonian Institu- tion Scholarly Press. Robbins, R. 2006. ?USAP Surface-Supplied Diving.? In Proceedings of the Advanced Scientifi c Diving Workshop, ed. M. A. Lang and N. E. Smith, pp. 187? 191. Washington, D.C.: Smithsonian Institution. Somers, L. H. 1988. ?Training Scientifi c Divers for Work in Cold Wa- ter and Polar Environments.? In Proceedings of Special Session on Coldwater Diving, ed. M. A. Lang and C. T. Mitchell, pp. 13? 28. Costa Mesa, Calif.: American Academy of Underwater Sciences. USAP. 1991. Standards for the Conduct of Scientifi c Diving. Arlington, Va.: National Science Foundation, Offi ce of Polar Programs. 17_Lang_pg241-252_Poles.indd 25217_Lang_pg241-252_Poles.indd 252 11/17/08 9:54:22 AM11/17/08 9:54:22 AM ABSTRACT. The under-ice environment places extreme selective pressures on polar ma- rine invertebrates (sea urchins, starfi sh, clams, worms) in terms of the low temperature, oligotrophic waters, and limited light availability. Free-swimming embryos and larvae face inordinate challenges of survival with almost nonexistent food supplies establishing near starvation conditions at the thermal limits of cellular stress that would appear to require large energy reserves to overcome. Yet, despite the long developmental periods for which these embryos and larvae are adrift in the water column, the coastal under-ice habitats of the polar regions support a surprising degree of vibrant marine life. How can so many animals be adapted to live in such an extreme environment? We all recognize that environmental adaptations are coded in the DNA sequences that comprise a species genome. The fi eld of polar molecular ecology attempts to unravel the specifi c imprint that adaptations to life in a polar habitat have left in the genes and genomes of these animals. This work requires a unique integration of both fi eld studies (under ice scuba diving and experiments) and laboratory work (genome sequencing and gene expression studies). Understanding the molecular mechanisms of cold adaptation is critical to our understanding of how these organisms will respond to potential future changes in their polar environments associated with global climate warming. INTRODUCTION: THE NECESSITY OF SCIENCE DIVING Looking across the coastal margins of most polar habitats, one is immedi- ately struck by the stark, frozen wasteland that hides the transition from land to sea beneath a thick layer of snow and ice. Standing on the sea ice surface along any shore line above 70? latitude, it is hard to imagine that there is fl uid ocean anywhere nearby, and even harder to think that there is even a remote possibility of animal life in such an environment. Yet under the fi ve meters of solid sea ice, a rich and active community of marine organisms exists. The real challenge is get- ting to them. Scientists studying how these organisms are adapted to survive and persist near the poles are limited by the logistical constraints in getting access under the ice to collect animals and plants for study. The sea ice cover establishes an effec- tive barrier to using most of the common collection techniques that marine biolo- gists employ from vessel-based sampling operations. The time and effort that is invested in opening a hole in the ice greatly precludes the number of sampling sites Adam G. Marsh, Professor, College of Marine and Earth Studies, University of Delaware, 700 Pilottown Road, Smith Laboratory 104, Lewes, DE 19958, USA (amarsh@udel.edu). Accepted 28 May 2008. Environmental and Molecular Mechanisms of Cold Adaptation in Polar Marine Invertebrates Adam G. Marsh 18_Marsh_pg253-264_Poles.indd 25318_Marsh_pg253-264_Poles.indd 253 11/17/08 9:22:32 AM11/17/08 9:22:32 AM 254 SMITHSONIAN AT THE POLES / MARSH that one can establish. Thus, from one ice hole, we need to be able to collect or observe as many animals as possible. To this end, scuba diving is an invaluable tool for the marine biologist because the ability to move away from the dive hole after entering the water greatly expands the effective sampling area that can be accessed from each individual hole. There is just no way to drill or blast a hole in the ice and drop a ?collection-device? down the hole and get more than one good sample. The fi rst deployment would collect what is under the hole, and after that, there is little left to collect. At present, there is still no better method (in terms of observational data, reliability, and cost effectiveness) than scuba diving for providing a scientist the necessary ac- cess to collect and study benthic marine invertebrates living along the coastal zones of polar seas. Although scuba diving under harsh, polar conditions is diffi cult and strenuous and not without risks, it is an absolute necessity for scientists to work in the water under the ice. We need to be able to collect, observe, manipulate, and study these unique polar marine invertebrates in their own environment. This is especially true for the develop- ing fi eld of environmental genomics, where scientists are attempting to decipher the molecular and genetic level changes in these organisms that make them so successful at surviving under extreme conditions of cold, dark and limited food. This paper will describe how important it is to ultimately understand how these adaptations work in the very sensitive early life stages of embryos and larvae, and how efforts to begin culturing these embryos and lar- vae under in situ conditions under the sea ice will contrib- ute to this greater understanding. ADAPTATIONS IN POLAR MARINE INVERTEBRATES Antarctic marine organisms have faced unique chal- lenges for survival and persistence in polar oceans and seas. The impacts of low temperatures and seasonally limited food availability have long been recognized as primary selective forces driving the adaptational pro- cesses that have led to the evolution of many endemic species in Antarctica (Clarke, 1991; Peck et al., 2004; Portner 2006; Clarke et al., 2007). Many elegant studies have demonstrated a wide-array of specifi c molecular ad- aptations that have been fi xed in specifi c genera or fami- lies of Antarctic fi sh, including chaperonins (Pucciarelli et al., 2006), heat shock proteins (Buckley et al., 2004; Hofmann et al., 2005), red blood cells (O?Brien and Sidell, 2000; Sidell and O?Brien, 2006), tubulin kinetics ( Detrich et al., 2000), and anti-freeze proteins (Devries and Cheng, 2000; Cheng et al., 2006). In contrast, the work with in- vertebrates has been less molecular and more focused on physiology and ecology, in terms of identifying adapta- tions in life-history strategies (Pearse et al., 1991; Poulin and Feral, 1996; Pearse and Lockhart, 2004; Peck et al., 2007), secondary metabolites ( McClintock et al., 2005), energy budgets and aging (Philipp et al., 2005a; Philipp et al., 2005b; Portner et al., 2005, Portner, 2006), and an ongoing controversy over the ATP costs of protein syn- thesis (Marsh et al., 2001b; Storch and Portner, 2003; Fraser et al., 2004; Fraser and Rogers, 2007; Pace and Manahan, 2007). Although much progress has been made over the last decade, we still barely have a glimmer as to the full set of adaptations at all levels of organization that have guided species evolution in polar environments. Each genetic de- scription, each physiological summary provides a snapshot of a component of the process, but we are still much in the dark as to how all the expressed phenotypes of an organ- ism are integrated into one functional whole, upon which selection is active. The survival of any one individual will depend upon the integrated effectiveness of ?millions? of phenotypic character states ranging from molecular to organismal level processes. How do you apply one snap shot to such a broad continuum spanning different orga- nizational scales? At present, there is a daunting lack of any specifi c indications of the ?key? genetic adaptations in single gene loci of any marine invertebrate. This is in stark contrast to the literature that exists for adaptations in fi sh genes and microbial genomes (Peck et al., 2005). Without the guidance of knowing ?where to look,? we are faced with identifying the best strategy for surveying an entire genome to pinpoint any possible genetic adaptations to survival in polar environments. The unique feature of cold-adaptation in polar marine invertebrates is that they are always exposed to a near- freezing temperature. Their entire lifecycle must be suc- cessfully completed at H110021.8?C (from embryo development to adult gametogenesis). Cold water (H110022 to 2?C) is an ex- treme environmental condition because of the complexity of the hydrogen bonding interactions between water mole- cules at the transition between liquid and solid phases. For a simple three atom molecule, the structure of water near freezing is very complex, with over four solid phases and the potential for two different liquid phases at low temper- atures. In polar marine invertebrates, signifi cant changes in the molecular activity of water at the liquid-solid inter- 18_Marsh_pg253-264_Poles.indd 25418_Marsh_pg253-264_Poles.indd 254 11/17/08 9:22:33 AM11/17/08 9:22:33 AM COLD ADAPTATION IN POLAR MARINE INVERTEBRATES 255 face (H110021.8?C) are likely to pose a strong selective force on biochemical and molecular function. Understanding how some organisms have adapted to this level of natural se- lection will provide information about the essential set of genetic components necessary for survival at low tempera- ture margins of our biosphere. EMBRYOS IN THE COLD The fact that polar marine invertebrates can main- tain a complex program of embryological development at low temperatures has received much discussion in the literature in terms of life-history adaptations. Of these ecological studies, Thorson?s rule has provided a focal point for numerous considerations of why development is so prolonged in marine invertebrates at high latitudes (Pearse et al., 1991; Pearse and Lockhart, 2004). The lim- ited availability of food in polar oceans has lead to un- certainty regarding the relative importance of low food availability compared with low temperatures as the pri- mary selective force limiting developmental rates (Clarke, 1991; Clarke et al., 2007). Overall, limits on metabolism have now received considerable attention in the literature and a general synthesis of the physiological constraints on organismal function in polar environments now appears to primarily involve cellular energetics at molecular and biochemical levels (Peck et al., 2004; Peck et al., 2006a; Peck et al., 2006b; Clarke et al., 2007). Recent work with the planktotrophic larvae of the Antarctic sea urchin, Ster- echinus neumayeri, has demonstrated that changes in the nutritional state of this feeding larvae do not alter its rate of early larval development (Marsh and Manahan, 1999; 2000). Low temperatures are likely a primary selective force and these larvae exhibit unique molecular adapta- tions to conserve cellular energy (Marsh et al., 2001b). This may be a general characteristic of polar invertebrate larvae, and could account for the predominance of non- feeding developmental modes found among benthic, mac- rofaunal invertebrates in Antarctica, particularly among echinoderms. EMBRYO ENERGY METABOLISM One of the most striking characteristics of embryonic development in Antarctic marine invertebrates is the slow rate of cell division. In the sea urchin S. neumayeri, early cleavage has a cell cycle period of 12 h, which is an order of magnitude slower than in a temperate sea urchin embryo at 15?C. In the Antarctic asteroid Odontaster validus the embryonic cell cycle period is just as long, and in the Ant- arctic mollusk Tritonia antarctica, it is extended to almost 48 h (Marsh, University of Delware, unpublished data). In general we assume that cell division is linked or coordi- nated to metabolic rates and that the increase in cell cycle period results from an overall decrease in metabolic rate processing at low temperatures. Embryos of S. neumayeri are sensitive to changes in temperatures around 0?C. Be- tween H110021.5?C and H110010.5?C, cell division exhibits a large change in the cycle period that is equivalent to a Q10 value of 6.2 (i.e., a 6.2-fold increase in cell division rates if ex- trapolated to a H900410?C temperature difference, Figure 1A). A Q10 greater than 3 indicates the long cell division cycles are determined by processes other than just the simple ki- netic effects of temperature on biochemical reaction rates. In contrast, metabolic rates in S. neumayeri do not evi- dence a change over this same temperature gradient (H90042?C; Figure 1B). There is no difference in oxygen consumption rates in embryos at the hatching blastula stage between H110010.5?C and H110021.5?C, despite large differences in cell divi- sion rates. This directly implies that the cell cycle period is not determined by a functional control or coordination to metabolic rates. The S. neumayeri results suggest that embryo development is not tied to metabolism at these low temperatures and the current notions of a selective mecha- nism that could favor patterns of protracted development by direct coordination to metabolic rates (ATP turnover) remain to be investigated. FIGURE 1. Temperature Q 10 for developmental rates and energy utilization in the Antarctic sea urchin Sterechinus neumayeri. (A) Embryogenesis at H110011.5?C was much faster than at ? 1.5?C and was equivalent to a 6-fold rate change per 10?C (i.e., Q10 estimate). (B) The metabolic rates of hatching blastulae at these two temperatures did not reveal any impact of temperature (i.e., Q10 H11011 0). 18_Marsh_pg253-264_Poles.indd 25518_Marsh_pg253-264_Poles.indd 255 11/17/08 9:22:33 AM11/17/08 9:22:33 AM 256 SMITHSONIAN AT THE POLES / MARSH EMBRYO MOLECULAR PHYSIOLOGY Some embryos of polar marine invertebrates appear to have specialized programs of gene expression that sug- gest a coordinated system of activity as a component of metabolic cold-adaptation. Most notable are the recent fi ndings in S. neumayeri embryos that mRNA synthesis rates are H110115-fold higher than in temperate urchin embryos (Marsh et al., 2001b). These data were determined from a time course study of whole-embryo RNA turnover rates and show that despite a large difference in environmental temperatures (H11011 H900425?C) rates of total RNA synthesis are nearly equivalent. In comparing the rate constants for the synthesis of the mRNA fraction there is a clear 5x up- regulation of transcriptional activity in the S. neumayeri embryos. What we need to know about this increased transcriptional activity is whether or not the upregulation of expression is limited to a discrete set of genes, or rep- resents a unilateral increase in expression of all genes. In order to perform these kinds of studies, we need to be able to work with embryos and larvae in their natural environ- ment under the sea ice. In addition to deciphering the magnitude of changes in transcriptional activities, we are now just beginning to realize the importance of how transcript levels may vary among individuals within a cohort. Variation is a necessity of biological systems. We generally think in terms of point mutations when we conceptualize the underlying basis of how individual organisms differ from one another within a species, and how novel phenotypes arise through the slow incremental accumulation of changes in nucleotide sequence (evolution). At odds with this ideology is the ob- servation that human and chimpanzee genes are too iden- tical in DNA sequence to account for the phenotypic dif- ferences between them. This lead A.C. Wilson (King and Wilson, 1975) to conclude that most phenotypic variation is derived from differences in gene expression rather than differences in gene sequence. Microarray studies are now revealing to us an inordinate amount of variation in gene expression patterns in natural populations, and we need to understand both the degree to which that variance may be determined by the environment, and the degree to which that variance may be signifi cantly adaptive. Although it is clear that interindividual variance in gene expression rates is a hallmark of adaptation and evolution in biological systems, at present, only a few studies have looked at this variation and the linkages to physiological function in fi eld populations. For Antarctic marine inverte- brates, most of the molecular and biochemical work look- ing at adaptations in developmental processes has focused on trying to fi nd ?extraordinary? physiological mecha- nisms to account for the adaptive success of these embryos and larvae. But what if the mechanism of adaptation is not extra-ordinary for polar environments? What if the mecha- nism is just ?ordinary? environmental adaptation: natural selection of individual genotype fi tness from a population distribution of expressed phenotypes. Understanding adap- tive processes in early life-history stages of polar marine in- vertebrates will ultimately require an understanding of the contribution that interindividual variance in gene expres- sion patterns plays in determining lifespan at an individual level, survival at a population level and adaptation at a spe- cies level in extreme environments. THE PROCESS OF ADAPTATION Natural selection operates at the level of an individ- ual to remove less-fi t phenotypes from subsequent gen- erations. However, it is clear that a hallmark of biological systems is the ?maintenance? of interindividual variance among individuals at both organismal (Eastman 2005) and molecular levels (Oleksiak et al., 2002; Oleksiak et al., 2005; Whitehead and Crawford, 2006). Although early life-history stages (embryos and larvae) are a very good system for looking at selective processes because there is a continual loss of genotypes/phenotypes during development, they are diffi cult to work with in terms of making individual measurements to describe a population (cohort) distribution. Their small size limits the amount of biomass per individual and consequently most of what we know about molecular and physiological processes in polar invertebrate larvae is derived from samples where hundreds to thousands of individuals have been pooled for a single measurement. However, methodological advances have allowed for quantitative measurements of molecular and physiologi- cal rate processes at the level of individual larvae in terms metabolic rates (Szela and Marsh, 2005), enzyme activities (Marsh et al., 2001a), and transcriptome profi ling (Marsh and Fielman, 2005). We are now beginning to understand the ecological importance of assessing the phenotypic vari- ance of characteristics likely experiencing high selective pressures. In Figure 2, the phenotype distributions of two species are presented to illustrate the signifi cant functional difference between how a change in the mean metabolic rate of a cohort (A) could be functionally equivalent to a change in the variance of metabolic rates within a cohort (B), where a decrease in metabolic rates (equivalent to an increase in potential larval lifespan) could arise from either 18_Marsh_pg253-264_Poles.indd 25618_Marsh_pg253-264_Poles.indd 256 11/17/08 9:22:35 AM11/17/08 9:22:35 AM COLD ADAPTATION IN POLAR MARINE INVERTEBRATES 257 process. Natural selection acts at the level of the phenotype of an individual, not at the level of the mean phenotype of a population or cohort. Thus, in order to understand most of the fi ne-scale biological processes by which organisms are adapted to polar environments (integrated phenotypes from multiple gene loci), we must be able to describe the distribution of the potential phenotypic space encoded by a genome relative to the distribution of successful (surviv- ing) phenotypes at a cohort (population) scale. Hofmann et al.?s recent review (2005) of the appli- cation of genomics based techniques to problems in ma- rine ecology clearly describes a new landscape of primary research in which it is possible to pursue the mechanis- tic linkages between an organism and its environment. One of the most interesting aspects of this revolution is the use of microarray hybridization studies for assessing gene expression profi les to identify the level of interindi- vidual variance that does exist in gene expression activities within a population (Oleksiak et al., 2005; Whitehead and Crawford, 2006). Although there are clearly some primary responses that organisms exhibit following specifi c envi- ronmental stresses or cues in terms of gene up- or down- regulation (Giaever et al., 2002; Huening et al., 2006), the power of assessing the expression patterns of thousands of genes simultaneously has opened an intriguing avenue of biological research: Why are gene expression patterns so variable, where is the source of that variation, what deter- mines the adaptive signifi cance of this variation? Although the mechanistic linkage between gene expression events and physiological rate changes may yet remain obscure, it is clear that a signifi cant level of biological variance is introduced at the transcriptome level, and the degree to which that variance may be signifi cantly adaptive requires exploration (Figure 2). HYPOTHESIS Overall, this research focus is attempting to describe a component of environmental adaptation as an integrative process to understand the mechanisms that may contrib- ute to long lifespans of larvae in polar environments. The over arching hypothesis is that embryos and larvae from the eggs of different females exhibit substantial variation in transcriptome expression patterns and consequently metabolic rates, and that these differences are an impor- tant determinant of the year-long survival of the few lar- vae that will successfully recruit to be juveniles. BIG PICTURE Variation is an inherent property of biological sys- tems. We know that genetic variation generates phenotypic variation within a population. However, we also know that there is more phenotypic variation evident within a population than can be accounted for by the underlying DNA sequence differences in genotypes. Much recent at- tention has been focused on the role of epigenetic infor- mation systems in regulating gene expression events, but there has been no consideration yet of the contribution of epigenetics to the level of variation in a phenotypic char- acter, whether morphological, physiological, or molecular. This idea focuses on a potential genome-wide control that could serve as a primary gating mechanism for setting limits on cellular energy utilization within a specifi c indi- vidual, while at the same time allowing for greater poten- tial interindividual variance in metabolic activities within a larval cohort. Understanding the sources of variation in gene transcription rates, metabolic energy utilization, and ultimately lifespans in polar larvae (i.e., the ?potential? range in responses to the selection pressures in polar envi- ronments) is essential for our understanding of how popu- lations are adapted to cold environments in the short-term, and ultimately how some endemic species have evolved in the long term. Within a cohort of larvae, it is now likely show that the variance at the level of gene expression events is FIGURE 2. Illustrated selection shifts in the frequency distributions of larval metabolic rates in an environment favoring greater energy conservation (black) over one that does not (gray). A change in met- abolic effi ciency within a cohort could arises at either the level of: (A) the mean phenotype or (B) the interindividual variance around the same mean in the phenotype. 18_Marsh_pg253-264_Poles.indd 25718_Marsh_pg253-264_Poles.indd 257 11/17/08 9:22:36 AM11/17/08 9:22:36 AM 258 SMITHSONIAN AT THE POLES / MARSH amplifi ed to a greater level of expressed phenotypic vari- ance. Thus, subtle changes in gene promoter methylation patterns (epigenetics) may have very pronounced impacts on downstream phenotypic processes (physiological en- ergy utilization). Describing how genomic information is ultimately expressed at a phenotypic level is vital for our understanding of the processes of organismal adaptation and species evolution. Although phenotypic plasticity is documented in marine organisms (particularly with re- gard to metabolic pathways), what is absolutely novel in this idea is that we may be able to demonstrate how changes in the phenotype distribution within a cohort of a character such as metabolic rates can routinely arise independently of genetic mutations (i.e., epigenetic con- trols). Conceptually, almost all studies of the regulation of gene expression events have been focused on a functional interpretation at the level of the fi tness of an individual. Our work to describe the variance in the distribution of expression rates within a group of larvae opens up a new dimension of trying to describe adaptational mechanisms at the larger level of total cohort fi tness. INTERINDIVIDUAL VARIATION IN EMBRYOS AND LARVAE DEVELOPMENT Embryos and larvae are rarely considered as popula- tions of individuals. They are normally just cultured in huge vats and mass sampled with the assumption that there is negligible interindividual variance among sibling cohorts. In S. neumayeri, we have observed substantial functional differences in the distribution of ?rate? pheno- types when the effort is made to collect data at the level of individual embryos and larvae. In Figure 3, eggs from fi ve females were fertilized and individual zygotes of each were scored for their rate of development to the morula stage (H110114 days at H110021.5?C). Even in this short span of time, there was clearly a difference in the embryo performance from different females with an almost two-fold variance in the mean cohort rates and an order of magnitude difference in the rates among all individuals. We normally think faster is better, at least in temperate and tropical marine environ- ments, but is that the case in polar environments? Are the fastest developing embryos and larvae the ones that are the most likely to survive and recruit 12 months later? ENERGY METABOLISM A novel methodology for measuring respiration rates in individual embryos and larvae has been developed for measuring metabolism in many individual embryos or lar- vae simultaneously (Szela and Marsh, 2005). This high- throughput, optode-based technique has been successfully used to measure individual respiration rates in small vol- umes (5 ul) as low as 10 pmol O 2 x h H110021 . The largest ad- vantage of this technique is that hundreds of individuals can be separately monitored for continuous oxygen con- sumption in real-time. The most striking observations we have made so far with S. neumayeri embryos is that there are large differences in metabolic rates between individu- als and that these differences appear to be infl uenced by the eggs produced by different females (Figure 4). Under- standing the distribution of metabolic rates among indi- viduals within a cohort of embryos or larvae is critical for understanding how metabolic rates may be ?tuned? to polar environments. In Figure 4, the most intriguing aspect of the distribu- tions is the 2- to 3-fold difference in metabolic rates that can be found among individual embryos. Clearly there is a large degree of interindividual variance in the cellular rate processes that set total embryo metabolism, and we need to understand the mechanistic determinants of that vari- ance. What we need to know now is how the biological variance at the level of the transcriptome (gene expression rates) impacts the variance at the level of physiological function (respiration rates). To date, the embryos and lar- vae of most polar invertebrates studied appear to evidence a strategy of metabolic down-regulation with the apparent FIGURE 3. Normal Distributions of development rates in late mor- ula embryos of S. neumayeri produced from different females. After fertilization, individual zygotes were transferred to 96-well plates and then scored individually for the time to reach 2-, 4-, 8-, 16-cell, and morula stages (n H11005 90 for each distribution). 18_Marsh_pg253-264_Poles.indd 25818_Marsh_pg253-264_Poles.indd 258 11/17/08 9:22:38 AM11/17/08 9:22:38 AM COLD ADAPTATION IN POLAR MARINE INVERTEBRATES 259 effect of extending larval lifespan (Peck et al., 2004; Peck et al., 2005; Peck et al., 2006a; Peck et al., 2006b). That is the observation at a population level. At an individual level this could be achieved by one of two mechanisms: (1) embryos in a cohort could maintain the same relative dis- tribution of metabolic rates around a lower mean rate (as in Figure 1A), or (2) embryos in a cohort could express a larger degree of interindividual variance (stochastic regu- lation) in respiration rates such that a larger fraction of the cohort would have lower metabolic rates that might concomitantly contribute to extended larval lifespans (as in Figure 1B). GENE EXPRESSION Physiological changes in metabolic rates must have an underlying basis in molecular events associated with gene expression rates. In order to compare changes within em- bryos or larvae to changes in gene activities, a novel meth- odology based on reannealing kinetics is employed for the rapid, high-throughput, effi cient, and economical profi ling of the sequence complexity of a transcriptome (Marsh and Fielman, 2005; Hoover et al., 2007a; b; c). Measuring re- naturation rates of cDNA along a temperature gradient can provide information about the transcript sequence complex- ity of a nucleic acid pool sample. In our assay, the reanneal- ing curves at discrete temperature intervals can be described by a second-order rate function. A full kinetic profi le can be constructed by analyzing all the curves using an informatics statistic (Shannon-Weaver entropy) for individual S. neu- mayeri embryos (hatching blastula stage). In Figure 5, each point represents the total mRNA pool complexity at a given Tm class with a mean (sd) of 4 duplicate assays at each Tm. Our novel revelation is that these mRNA pools among em- bryos are not identical. We can detect discrete differences in the distribution and abundance of component transcripts at the level of these individuals, even though they are all apparently progressing through the same developmental program. Thus, there is a large variance component that arises at the level of gene expression within these Antarctic sea urchin embryos. One of the most prominent mechanisms determining gene expression patterns is the system of chemical modifi - cations on DNA that establish an epigenetic pattern of in- formation. Epigenetic processes refer to heritable mole cular structures that regulate gene expression events independent of any DNA nucleotide sequence within a genome. There is currently a clear recognition that a large component of gene regulation can operate at this level of local DNA structure and composition and that DNA methylation is one of the dominant mechanisms. Methylation of promoter domains is one of the key mechanisms that can account for temporal patterns of differential gene expression during embryologi- cal development (MacKay et al., 2007; Sasaki et al., 2005; Haaf, 2006). During cell division, this genome methylation pattern is re-established with high fi delity in daughter cells by a suite of methyl-transferases immediately after DNA synthesis. Consequently, an established regulatory ?im- print? can be perpetuated during development and across generations. In this sense, methylation serves as a cell-based FIGURE 5. Individual cDNA libraries were constructed for four em- bryos and profi led for sequence complexity using a novel approach to measuring reannealing kinetics. Large differences in the sequence distribution and abundance of the transcriptome pool are evident among these sibling embryos. FIGURE 4. Distribution of respiration rates in late gastrula embryos of S. neumayeri. Eggs from 6 females were fertilized with the sperm from one male and maintained as separate cultures. The distribution of metabolic rates in these cultures is not equivalent (n H11005 15 for each culture; 90 individuals total). 18_Marsh_pg253-264_Poles.indd 25918_Marsh_pg253-264_Poles.indd 259 11/17/08 9:22:39 AM11/17/08 9:22:39 AM 260 SMITHSONIAN AT THE POLES / MARSH memory system for maintaining a pattern of gene expres- sion regulation. Methylation is also evident in invertebrate genomes, although the distribution of methylated sites appears to be more variable than in vertebrates, being mainly con- centrated within areas of gene loci (Levenson and Sweatt, 2006; Schaefer and Lyko, 2007; Suzuki et al., 2007). Some invertebrates exhibit mammalian-type levels of DNA meth- ylation (up to 15 percent of all cytosine residues methyl- ated). In many invertebrates, DNA methylases appear to be very active during development (Meehan et al., 2005) and we can measure H110114 percent methyl-cytosine levels in S. neumayeri (Kendall et al., 2005). More importantly, the presence/absence of specifi c methylation sites within a ge- nome can be rapidly assessed using an Amplifi ed Fragment Length Polymorphism derived assay. The key observations here are that methylation fi ngerprints can be rapidly as- sessed following experimental treatments, and that meth- ylated sites in gene promoters can be identifi ed and scored separately (Kaminsk et al., 2006; Rauch et al. 2007). The number and diversity of methylation sites we can identify in early S. neumayeri embryos indicates an active DNA methylation system that could be generating the variance in metabolic rates and developmental rates that we have observed. DIVING WITH EMBRYOS AND LARVAE UNDER THE ICE The previous sections have documented the prevalence of a high component of interindividual variance among individual embryos of S. neumayeri. In order to fully un- derstand the implications for this variance in terms of its impact on the survival of a cohort of larvae and the per- sistence of a sea urchin population through time, we need to study lots of individual larvae under natural conditions. This is absolutely impossible to do in a laboratory setting because of the large volumes of sea water that would have to be maintained. As an alternative, a pilot program is underway to investigate the success of culturing embryos and larvae in fl ow-through containers under the sea ice in McMurdo Sound. The concept is simple: Let the embryos and larvae grow naturally without investing much time or effort in their husbandry. Current efforts to mass culture sea urchin larvae uti- lize 200-liter drums stocked at densities of H1101110 per ml. At those densities, the water needs to be changed every two days. One water change on one 200 liter drum can take 2 hours in order to ?gently? fi lter all the larvae out fi rst and then put them into another clean 200 liter drum. The word gently is emphasized here for irony, because there is nothing gentle about using a small-mesh screen to fi l- ter out all the embryos and larvae. It is a very physically stressful process, and mortality rates are signifi cant. An alternative approach utilizing in situ chambers could allow for the embryos and larvae to develop without human intervention. The under ice environment around McMurdo Station is very stable and the epiphytic foul- ing community of organisms is almost nonexistent. In Figure 6, photographs of under ice culture bags in use in McMurdo Sound are shown. These trial bags were fi tted with two open ports having a 40 H9262m mesh Nytek screen coverings so that water could be freely exchanged through the bag. Low densities of embryos from different marine FIGURE 6. Culture bags with embryos and larvae of different ma- rine invertebrates are tethered to the bottom rubble of the McMurdo Jetty under 6 m of solid sea ice and 25 m total water depth (photos by Adam G. Marsh). 18_Marsh_pg253-264_Poles.indd 26018_Marsh_pg253-264_Poles.indd 260 11/17/08 9:22:41 AM11/17/08 9:22:41 AM COLD ADAPTATION IN POLAR MARINE INVERTEBRATES 261 invertebrates were placed in the bag and sampled at dif- ferent time intervals. We have been successful at culturing larvae of the clam Laternula eliptica for over 13 months under the ice, and recovering fully metamorphosed juve- niles at the end of that time period. The ability to place embryos and larvae under the ice in culture containers in the Austral summer, then leave them in place through the winter season, means that we now have the potential to work with the full lifecycle of some marine invertebrate larvae. Because of the slow de- velopmental rates of these larvae and protracted lifespans, there are relatively few studies that have been able to col- lect any data on the later life-stage as they approach meta- morphosis. However, divers can now establish cultures in situ under the ice, then return at any time point later on to sample individuals. This increase in the time span for which we can now study will fi ll in the large gap in our current knowledge of what happens at the end of the Aus- tral winter period when the larvae are ready to become juveniles. In order to understand how larvae are adapted to survive in polar environments, we really need to make our best measurements and execute our most exact ex- periments on the individuals that have survived for the complete developmental period and are now ready to be- come juveniles. If only 10 percent of a cohort survives to this stage (H1100112 months development), then making early measurements on the other 90% (at 1 month) that were destined to die would essentially just give you information about what does not work. It is the survivors that hold the key to understanding how these organisms are adapted to persist in a harsh polar environment. In essence, all the existing studies on adaptations in polar marine larvae that have made measurements from bulk cultures at early de- velopmental time points could be grossly misleading. All of those individuals are not likely to survive the full year to recruitment. Consequently, we need an experimental cul- turing system that will allow scientists to work with larvae that have survived the harsh polar environment. Those are the individuals that have the key to understanding adap- tive processes. Overall, the in situ culturing approach offers us three main advantages: 1. Large numbers of individuals can be cultured with rela- tively little husbandry effort. Once the culture contain- ers are setup and stocked, then the only effort necessary is for a dive team to periodically sample and remove individuals. There is no feeding and little maintenance required. 2. Larvae can be cultured under very natural conditions without laboratory artifacts. The most important vari- able to control is temperature and by not having open culture containers in an aquarium room, there is no worry about the temperature in the vessels changing because of problems with electricity supply, pumps breaking down and losing the cold sea water supply rate, or someone just changing the thermostat within the aquarium room. The second most important vari- able is food, and under in situ conditions, the feeding larvae will receive a diet of natural species and in a natural supply. 3. Long term cultures can be maintained across the entire developmental period, which in most polar marine in- vertebrates can easily extend upwards of a year. This approach will provide access to larvae that have suc- cessfully survived their full lifecycle in a harsh polar environment. CONCLUSION The S. neumayeri distributions in developmental rate (Figure 3), respiration distributions (Figure 4), transcrip- tome profi les (Figure 5), and gene methylation (data not shown) have focused our attention on trying to under- stand the functional signifi cance of interindividual vari- ability at these levels of biological organization. In a life- history model that selects for a prolonged larval lifespan, it is intriguing to ask whether or not it is a reduction in individual metabolic rates (Figure 2A) or an increase in the cohort variance in metabolic rates (Figure 2B) that could account for the adaptation in metabolic phenotypes. The metabolic lifespans of polar invertebrate larvae could be under the same genetic determinants as other temperate species, but changes in patterns of gene regulation could substantially alter the distribution of physiological pheno- types within a cohort. Being able to study long-lived larvae that are ready to become juveniles holds the key for deci- phering the adaptive mechanism that may be operative at the level of a full cohort to ensure that some percentage is capable of surviving. Selection is surely not operating to force the survival function of all individuals within a cohort. Only enough need to survive to keep a population established and stable. Scientifi c diving will be an important component of discovering how these animals are adapted to survive. The opportunity to now work in situ with embryos and larvae will open new avenues of research and understanding. Even though the under ice work is not complex, it is nonetheless 18_Marsh_pg253-264_Poles.indd 26118_Marsh_pg253-264_Poles.indd 261 11/17/08 9:22:45 AM11/17/08 9:22:45 AM 262 SMITHSONIAN AT THE POLES / MARSH rigorous and demanding. Any diver working on the bottom for more than 45 minutes will readily attest to the extreme nature of the cold that impacts all organisms in that envi- ronment. Being in the water gives one a unique perspective on the survival challenges that are facing the embryos and larvae of these polar marine invertebrates. ACKNOWLEDGMENTS This work was conducted with the help, collaboration, and participation of numerous students and postdoctoral associates in my lab and I am grateful for their efforts, energies and cold tolerance. The research was supported by a grant from the National Science Foundation, Offi ce of Polar Programs (#02? 38281 to AGM). LITERATURE CITED Buckley, B. A., S. P. Place, and G. E. Hofmann. 2004. 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Physiological and Biochemical Zoology, 76: 28? 40. Suzuki, M. M., A. R. W. Kerr, D. DeSousa, and A. Bird. 2007. CpG Methylation is Targeted to Transcription Units in an Invertebrate Genome. Genome Research, 17: 625? 631. Szela, T. L. and A. G. Marsh. 2005. Microtiter Plate, Optrode Respi- rometry Reveals Large Interindividual Variance in Metabolic Rates among Individual Nauplii of Artemia sp. Marine Ecology Progress Series, 296: 291? 309. Whitehead, A., and D. L. Crawford. 2006. Variation within and among Species in Gene Expression: Raw Material for Evolution. Molecular Ecology, 15: 1197? 1211. 18_Marsh_pg253-264_Poles.indd 26318_Marsh_pg253-264_Poles.indd 263 11/17/08 9:22:46 AM11/17/08 9:22:46 AM 18_Marsh_pg253-264_Poles.indd 26418_Marsh_pg253-264_Poles.indd 264 11/17/08 9:22:46 AM11/17/08 9:22:46 AM ABSTRACT. McMurdo Sound, Antarctica, is the best place to conduct diving physiol- ogy studies on marine birds and mammals under free-diving conditions. Both emperor penguins and Weddell seals live naturally in areas of extensive sea ice under which they dive and hunt for prey. The fi rst experimental diving studies were conducted on Wed- dell seals in 1964 using the isolated breathing hole protocol for the fi rst time. Sea ice, 2 m thick, covers McMurdo Sound until late December. Below the ice is the deepwater environment where Antarctic predators hunt their prey. Here in the Sound diving studies involve attachment of a recording device to a seal or bird and release of the animal into the hole cut in sea ice. This procedure sets the stage for a bird or mammal to hunt with- out competition, and the only restrictive condition is that they must return to the release hole to breathe. After the animal surfaces, the attached recording devices can be retrieved and the information downloaded. Results from using this experimental protocol range from determining the fi rst foraging patterns of any diving mammal, to measuring the fi rst blood and muscle chemistry fl uctuations during the extended and unrestrained dives. These experiments are the standard for understanding the hypoxic tolerance of diving animals, their aerobic diving limits, and their strategies of foraging, to mention a few. The protocol will continue to be used in 2008 for studies of both emperor penguins and Weddell seals by several investigators. INTRODUCTION My goal in this presentation is to engender an understanding of the valu- able resource we have next to McMurdo Station, the largest base in Antarctica. That asset is McMurdo Sound itself, which is covered by perhaps the largest and most southerly annual fast-ice sheet in Antarctica. The ice cover most commonly ranges in thickness from 1 to 4 m in thickness, and extends from the McMurdo Ice Shelf to Cape Royds to the north, and east to west from Ross Island to the continent (Figure 1). It covers an oceanic area reaching to depths of 600 m. It is also one of the most stable fast ice areas and it persists until late December. Like almost every fi rst-time visitor, when I arrived in 1961, I was not impressed by the Sound?s uniqueness. However, McMurdo Sound may have the only such annual fast-ice shelf anywhere in Antarctica where it can be used extensively for innumerable projects. Among the many uses are: (1) the largest and most active airport in Antarctica, (2) ?at-sea? marine biological and oceanographic stations Gerald Kooyman, Research Professor, Scholander Hall, Scripps Institution of Oceanography, Univer- sity of California? San Diego, 9500 Gilman Drive, La Jolla, CA 92093-0204, USA ( gkooyman@ ucsd.edu). Accepted 28 May 2008. Milestones in the Study of Diving Physiology: Antarctic Emperor Penguins and Weddell Seals Gerald Kooyman 19_Kooyman_pg265-270_Poles.indd 26519_Kooyman_pg265-270_Poles.indd 265 11/17/08 9:29:57 AM11/17/08 9:29:57 AM 266 SMITHSONIAN AT THE POLES / KOOYMAN without the inconveniences and cost of research vessels, (3) scuba diving stations, and (4) at least three to four ex- perimental laboratories for the study of marine organisms. These stations are scattered throughout the Sound in the spring and sometimes in the winter. As an example of the Sound?s value as a scientifi c asset I will describe what is most familiar to me. For the past 43 years, the Sound has been the premier study site for the investigation of diving and behavioral physiology of birds and mammals, and the training of three generations of scientists. These kinds of studies began not long after the station was established in 1957, and there was a surge in scientifi c endeavor pro- moted by, and in celebration of the Second International Polar Year, or International Geophysical Year (IGY), as it was called then. The diving studies have been continuous ever since. The crucial attributes that a plate of fast ice must have to make it useful year-round, is a large surface area of at least 10s of km 2 . It must have an ice thickness that will support large vehicles such as Caterpillar D8?s, substan- tial buildings, and large aircraft of at least the size of an LC 130. McMurdo Sound fast ice will support the Boeing C5, the largest aircraft known. Ross Island, where Mc- Murdo Station resides, also provides protection for the Sound from ocean currents and storms so that the annual fast ice persists until early January. In fact, in the last few years, since about 2001 until 2006, little of the Sound ice broke up and departed. This was a result of the added protection given to the Sound by the giant iceberg B15. As a consequence of the Sound?s fast-ice stability, the largest airport in Antarctica was built in McMurdo Sound dur- ing the IGY (1957), and a new one has been built every spring since that fi rst season. In addition, there is a large, local Weddell seal population in McMurdo Sound that has been the object of intensive ecology and physiology stud- ies since the establishment of the two bases of McMurdo Station and Scott Base. McMurdo Sound fulfi lls all the above requirements. It is about 64 km across the Sound from Ross Island to the mainland, and it is about 32 km from Cape Royds to the McMurdo Ice Shelf near Cape Armitage (Figure 1). The sea ice forms in April and decays in January, usually breaking out annually on the eastern half by mid to late February. The ice grows through October at which the maximum is usually about 2 m thick at its southern base next to the McMurdo Ice Shelf, and about 1.5 m thick near the edge at Cape Royds. It is noteworthy to illustrate the variability of the sea ice breakout, formation, and extent. In 1981 during the fi rst overwintering study of Weddell seals, investigators were hampered by the late development of sea ice well into the winter after a previous extensive summer ice breakout to the McMurdo Ice Shelf. In contrast, in 2001 after the ar- rival of B15 at the northern edge of Ross Island, the annual fast ice became multi-year ice and extended well beyond Beaufort Island. This condition persisted until 2006 when the last remnants of this iceberg drifted north of the zone of infl uence on McMurdo Sound. Because of the numerous science programs at McMurdo Station and Scott Base, as well as McMurdo Station?s function as the logistic center for supplying South Pole Station, the airport, that has hun- dreds of landings every season, is essential for this region of Antarctica. In addition, sea ice to land access for large vehicles to reach McMurdo Station and Scott Base is ideal with gently sloping land down to the sea ice edge. Finally, the Weddell seal population along the coast of Ross Island from Turtle Rock to Cape Royds (~15 km) harbors about 500 breeding females and it is one of the largest concentra- tions of seals in the Ross Sea. The above-described attributes of McMurdo Sound are matchless. There are no other stations throughout Ant- arctica that have the air support or base size and support of McMurdo Station. Consider the rest of the Ross Sea. There are two possibilities: Terra Nova Bay (TNB) and Moubray Bay both of which have extensive fast ice sheets. In the southern end of Terra Nova Bay resides the Italian base of Zuchelli Station. The Campbell Ice Tongue bisects the bay. The small southern section would be feasible for only limited bird and mammal work. Here there is a small FIGURE 1. Location of McMurdo Sound. The annual sea ice north- ern limit is usually at Cape Royds, but occasionally extends to Cape Bird. The two major research stations of McMurdo Station (US) and Scott Base (NZ) are near the tip of Cape Armitage. 19_Kooyman_pg265-270_Poles.indd 26619_Kooyman_pg265-270_Poles.indd 266 11/17/08 9:29:58 AM11/17/08 9:29:58 AM MILESTONES IN THE STUDY OF DIVING PHYSIOLOGY 267 airstrip and a few offshore marine stations. The northern portion of TNB is much larger and has both a large popu- lation of Weddell seals, and one of the largest known em- peror penguin colonies. This part of the bay is bordered by high ice cliffs and is not accessible from land. There is also a large perennial ice crack that bisects the eastern part of TNB from east to west that limits its usefulness for bird and mammal studies. There is also no access to land so that a shore station could not be established to support the kind of programs that are carried out from McMurdo Station and, at least for seals, it would be diffi cult to establish a func- tional sea ice station where the isolated hole protocol (IHP) would work. Wood Bay to the north of Cape Washington has no bases except for a small fi eld camp at Edmonston Point for Ad?lie penguin research, and there is no good site for a station. Little is known about this area, but it appears to have a substantial seal population. The only accessible land to sea ice in Moubray Bay is on Hallett Peninsula where a large Ad?lie penguin colony occupies the entire land surface area, and consequently is not an option for a research station now, although there was a research station there in the past. In the eastern Ross Sea adjacent to Cape Colbeck, Bartlett Inlet forms about a 15 km bight into an embayment of fast ice, which at the most southern extension is found a large emperor penguin colony. Little is known about this very isolated region that is notorious for foul weather and extensive pack ice. There is no easy access to land where a permanent station might be established. All of the described areas are small com- pared to McMurdo Sound and would provide a much more limited program of research than is possible at McMurdo Sound. These are the areas that I have fi rst hand knowledge. As far as the rest of Antarctica is concerned, to my knowl- edge, there are no areas with air strips to handle routine air support from outside of Antarctica, with the exception of the Antarctic Peninsula, and none of those in the peninsula have extensive fast-ice sounds for the conduct of marine re- search. In short, there are no other places in the Antarctica or the world, where marine research, especially on birds and mammals can be conducted in the way I describe below. METHODS With a 2-meter-thick layer of ice over the Sound, the fi rst objective is to pierce through it to reach the marine environment below. In the early days, this was a major, backbreaking task to cut a hole through the ice with a chainsaw and lift out the cut blocks from the developing hole with ice tongs. Depending upon the number of labor- ers and the thickness of the ice, the task of penetration could take from a few hours to several days. Thankfully, soon after the cutting task was assisted by using explo- sives and the sea ice landscape began to be peppered with marine stations. However, this was not ideal for several reasons one of which was the potential harm to seals div- ing under the ice. By the early 1970s, a 1.2 m diameter augur was employed for hole cutting. Because the holes even in the thickest ice can be cut in about 20 min, this has become one of the most valuable assets in the toolbox of marine work in McMurdo Sound. The sea ice stations have a range of functions. Some are platforms for setting fi sh and invertebrate traps to cap- ture benthic specimens for ecology and physiology stud- ies. Others are used for setting up a long-line to capture the large Antarctic cod, Dissostichus mawsoni, which may weigh greater than 100 kg. From these specimens much has been learned about anti-freeze properties of the body fl uids, their physiology, and their natural history. My own personal experience has been to use the sea ice stations for conducting detailed diving studies on unrestrained Weddell seals and emperor penguins. For these kinds of experiments there is no match for the situation. In 1964 I established the isolated hole protocol wherein the dive hole was established a distance of 1 to 4 km from any other hole, either manmade or natural. Under these cir- cumstances a penguin or seal released into the isolated hole had to return to this same breathing hole, and there- fore, instruments could be deployed and recovered after each or a series of subsequent dives. Normally the diving seals would remain and use the hole for hours to days. During this period the hole was covered with a heated hut for the convenience of the investigators. The emperor pen- guins were peculiar because they would never surface in a hole covered with a hut if they had an option to go to a hole in the open. At the open air hole they would leave the water after almost every dive. The IHP has been used continuously by a variety of investigators over the past 43 years (as of 2007), and several projects are planned for the future. A great deal of information has been collected during these studies and the following are a few highlights most familiar to me. RESULTS (BIRDS AND MAMMALS) BEHAVIOR? WEDDELL SEALS The fi rst diving studies ever conducted on a diving an- imal, in which detailed diving information was obtained, occurred in 1964 in McMurdo Sound. Using time-depth 19_Kooyman_pg265-270_Poles.indd 26719_Kooyman_pg265-270_Poles.indd 267 11/17/08 9:30:00 AM11/17/08 9:30:00 AM 268 SMITHSONIAN AT THE POLES / KOOYMAN recorders full advantage of the IHP was used to determine the behavior and physiology of Weddell seals (Kooyman, 1968). Investigations of this kind have been in progress ever since. The fi rst results broke new ground in many ways, and one of the most signifi cant was to show that we had been far too conservative in assumptions about marine mammal diving capacities. Indeed, even the fi rst two publications on Weddell seals were too conservative on the estimates of what these animals could do (De Vries and Wohlschlag, 1964; Kooyman, 1966). In the latter re- port it was proposed that the maximum depths and dura- tions were proposed to be 600 m and 46 min, respectively. At present the depth and duration records for Weddell seals now stand at 714 m (Testa, 1994), and 96 min (Za- pol, Harvard Medical School, personal communication). Seals accomplished all of these exceptionally long dives while diving from an isolated hole. This procedure brings out the extremes in breath holding of these animals. Pre- sumably they are responding to the trauma of capture and transport to the hole, and are trying to fi nd an escape route from the new environment. However, within a few hours they settle in to routine hunting dives, and take advantage of the isolation and being away from competi- tion with other seals. This provides to the investigator the best of both worlds. One is the discovery of some of the limits the seals may press themselves toward followed by the ordinary kind of effort that they do routinely. Both are of interest to the behaviorist and physiologist. BEHAVIOR? EMPEROR PENGUINS Emperor penguins have responded in a somewhat dif- ferent way from Weddell seals to the IHP. They still achieve some of the longest dives recorded by emperor penguins and this record stands at 23 min (P. J. Ponganis, Scripps Institution of Oceanography, personal communication) compared to 21.8 min obtained from a free ranging bird (Wienecke et al., 2007). However, emperor penguins sel- dom make deep dives during the IHP compared to animals foraging under more natural conditions of the pack ice. The maximum depth of the IHP is 250 m (Ponganis et al., 2004), while that of the free ranging animals is 560 m (Wienecke et al., 2007). At least part of the reason for the shallow dives by emperor penguins during IHP is the lack of incentive. Free ranging birds feed primarily on Antarc- tic silver fi sh at mid-water depths in the Ross Sea. How- ever, birds hunting under thick fast ice of the Sound switch from silver fi sh, even though these are abundant and the primary prey of Weddell seals, to another fi sh, Pagothenia borchgrevinki, which is present in large numbers just un- der the ice. This has been a frustration to the physiologist who wants to explore the responses of the animals under extreme conditions. However, deep dives are so rare that it is purely by chance when physiological protocols are in place when the animals dive to extremes. Indeed, in the many thousands of dives observed in Weddell seals only once were some of the responses made. In one case, a blood lactate sample was obtained after a 66 min dive (Kooyman et al., 1980), and pulmonary function measurements were obtained from a seal that made an 82 min dive (Kooyman et al., 1971). However, a benefi t for the behaviorist and ecologist working in McMurdo Sound is that it has been possible to deploy ?Crittercams? or ?VDAPS? on emperor penguins and Weddell seals, respec- tively. These are animal borne imaging devices that have made it possible to observe their hunting tactics. With the Crittercam on emperor penguins taped images show how the birds catch P. borchgrevinki hiding in ice crystals at- tached to the fast ice undersurface (Ponganis et al., 2000). Similarly, with the VDAP Weddell seals? captures of Ant- arctic silverfi sh and Antarctic cod were documented (Davis et al., 1999). All of these imaging results proved exciting both to the scientifi c community and the media. PHYSIOLOGY? WEDDELL SEALS For the comparative physiologist working on diving physiology McMurdo Sound has been a magnet because of the IHP. This protocol has made possible the attach- ment of complex instruments to record heart rate, measure blood chemistry, and to determine oxygen and nitrogen tension in blood at known depths and times in the dive. Some of the results have been the generation of the lactate endurance curve from blood samplings of several differ- ent seals (Kooyman et al., 1980). This curve has defi ned the aerobic diving limit (ADL) of Weddell seals by the in- fl ection area within the curve (Figure 2). The concept was defi ned by Kooyman (1985), and states that ?The ADL is defi ned as the maximum breath hold that is possible without any increase in blood LA concentration during or after the dive.? It was further explained that the calculated ADL (cADL) could be estimated if the body oxygen store and diving metabolic rate were known. This concept has motivated many research projects to make estimates of the ADL in a variety of diving animals. Several of these studies have been done on Antarctic divers including the emperor penguin using the IHP for the study in McMurdo Sound. Another landmark study on Weddell seals, among the many that have occurred, was the determination of blood N 2 levels while Weddell seals were diving to depth 19_Kooyman_pg265-270_Poles.indd 26819_Kooyman_pg265-270_Poles.indd 268 11/17/08 9:30:01 AM11/17/08 9:30:01 AM MILESTONES IN THE STUDY OF DIVING PHYSIOLOGY 269 (Falke et al., 1985) (Figure 3). This result corroborated the more artifi cial studies conducted on elephant seals forc- ibly submerged and compressed in a hyperbaric chamber ( Kooyman et al., 1971) a number of years earlier. The sa- lient feature of these blood pN 2 results was that no matter the depth of the dive the N 2 tensions do not rise above a relatively low value that is unlikely to cause decompres- sion sickness. The only similar measurements on other species diving under unrestrained conditions are those of the emperor penguin using the IHP in McMurdo Sound (Ponganis, personal communication). PHYSIOLOGY? EMPEROR PENGUINS Using the same procedures as applied to the Weddell seal, the ADL of emperor penguins has been determined (Ponganis et al., 1997), and it has equally important im- plications for diving birds as the studies of Weddell seals had for diving mammals. To probe this problem further a recent study has asked the question of how emperor pen- guins manage their oxygen stores. This question is of ex- ceptional importance to understanding the physiological principles of how diving animals overcome the problems of limited oxygen and endure hypoxia on a routine basis. In the course of these studies the investigators have shown that emperor penguins can tolerate exceptionally low ar- terial O 2 tensions in the range of 5? 10 mmHg (Ponganis, personal communication). These values are at levels be- low what could maintain consciousness in most terrestrial birds and mammals, including humans. Indeed they are substantially below the expected arterial O 2 tension of a person standing on Mt. Everest (Chomolungma), and they raise a series of questions about how this diving bird over- comes such extreme hypoxia. DISCUSSION I have briefl y mentioned several types of studies on birds and mammals that have had a signifi cant impact on diving behavior and physiology of marine animals. If this were a comprehensive review of work accomplished in McMurdo Sound there would be many disciplines and investigators represented and the number of publica- tions would be in the hundreds if not thousands. Many of the studies could not have been done anywhere other than in McMurdo Sound. As for the birds and mammals studies there are at least three generations of investigators, FIGURE 2. Lactate endurance curve of the Weddell seal. The gray dots are peak arterial post-dive lactic acid concentrations of aero- bic dives. The black dots are the peak arterial post-dive lactic acid concentrations after dives with a lactate accumulation above resting levels (modifi ed from Kooyman et al., 1980). FIGURE 3. Arterial N 2 tensions in a Weddell seal. The seal made a free dive to 89 m under McMurdo Sound fast ice. The dive began at zero on the abscissa, reached a maximum depth at 5 min, and the dive ended after 8 min. 19_Kooyman_pg265-270_Poles.indd 26919_Kooyman_pg265-270_Poles.indd 269 11/17/08 9:30:01 AM11/17/08 9:30:01 AM 270 SMITHSONIAN AT THE POLES / KOOYMAN spanning 45 years that have found the greatest contribu- tions to their fi eld in McMurdo Sound. In regard to the diving studies mentioned above, many of the results are fundamental to the fi eld of behavior and physiology of diving. Refl ecting on these accomplishments and many others such as the long-term works of Siniff?s trends in Weddell seal populations (Siniff et al., 1977), DeVries?s range of work on ecology to the molecular nature of antifreeze com- pounds in fi sh (DeVries, 1971) and long term catch data on the highly sought after Antarctic cod (DeVries, Univer- sity of Illinois, unpublished data), and Dayton?s cage ex- clusion studies of benthic organisms (Dayton et al., 1974; Dayton, 1985), I wonder if McMurdo Sound is not under- appreciated. In the early days it was not coveted at all, and was used as a dumping ground for human waste and many harmful inorganic and organic products. Thankfully those ?bad old days? ended a long time ago, and like Antarctica in general, it is treated by a much softer human footprint. Still, to recognize the distinct value of McMurdo Sound it should be given some sanctuary or museum status as the great contributor to biological sciences, because as I hoped to achieve in this review, there is no other place like it. We do not know what specifi c effects the changing climate in Antarctica will have on the seas that surround the continent, but with the long-term history of science in McMurdo Sound, it will be one of the areas invaluable in assessing some of the changes. Especially with the large modern laboratory of Crary Center adjacent to the Sound where researchers can study the animals with some of the most sophisticated techniques in the world. ACKNOWLEDGMENTS Thanks to Smithsonian for making this symposium possible and providing the travel funds to attend. All my work over the past 45 years at McMurdo Sound has been supported by numerous NSF OPP grants for which I will always be grateful. I give thanks, as well, to the many support contractors and their staff on land, sea, and air that have made the journeys such a fi ne experience. In the background always has been my wife Melba and my sons, Carsten and Tory. It was also an inestimable joy to see both sons become the best fi eld assistants one could ever want. LITERATURE CITED Davis, R., L. Fuiman, T. Williams, S. Collier, W. Hagey, S. Kanatous, S. Kohin, and M. Horning. 1999. Hunting Behavior of a Marine Mammal beneath the Antarctic Fast Ice. Science, 283: 993? 996. Dayton, P. K. 1985. Antarctica and Its Biota (A Review of Antarctic Ecology). Science, 229: 157? 158. Dayton, P. K., G. A. Robilliard, R. T. Paine, and L. B. Dayton. 1974. Bio- logical Accommodation in the Benthic Community at McMurdo Sound, Antarctica. Ecology (Monograph), 44: 105? 128. DeVries, A. L. 1971. Glycoproteins as Biological Antifreeze Agents in Antarctic Fishes. Science, 172: 1152? 1155. DeVries, A. L., and D. E. Wohlschlag. 1964. Diving Depths of the Weddell Seal. Science, 145: 292. Falke, K. J., R. D. Hill, J. Qvist, R. C. Schneider, M.Guppy, G. C. Liggins, P. W. Hochachka, and Z. Elliot. 1985. Seal Lungs Collapse during Free Diving: Evidence from Arterial Nitrogen Tensions. Science, 229: 556? 558. Kooyman, G. L. 1966. Maximum Diving Capacities of the Weddell Seal (Leptonychotes weddelli). Science, 151: 1553? 1554. Kooyman, G. L. 1968. An Analysis of Some Behavioral and Physiologi- cal Characteristics Related to Diving in the Weddell Seal. In Biol- ogy in the Antarctic Seas III, ed. G. A. Llano and W. L. Schmitt. Antarctic Research Series, Vol. 11. Washington, D.C.: American Geophysical Union. ???. 1985. Physiology without Restraint in Diving Mammals. Marine Mammal Science, 1: 166? 178. Kooyman, G. L., D. H. Kerem, W. B. Campbell, and J. J. Wright. 1971. Pulmonary Function in Freely Diving Weddell Seals (Leptonychotes weddelli). Respiratory Physiology, 12: 271? 282. Kooyman, G. L., E. Wahrenbrock, M. Castellini, R. W. Davis, and E. Sinnett. 1980. Aerobic and Anaerobic Metabolism during Volun- tary Diving in Weddell Seals: Evidence of Preferred Pathways from Blood Chemistry and Behavior. Journal of Comparative Physiology, 138: 335? 346. Ponganis, P. J., G. L. Kooyman, L. N. Starke, C. A. Kooyman, and T. G. Kooyman. 1997. Post-Dive Blood Lactate Concentrations in Em- peror Penguins, Aptenodytes forsteri. Journal of Experimental Biol- ogy, 200: 1623? 1626. Ponganis, P. J., R. P. Van Dam, G. Marshall, T. Knower, and D. Levenson. 2000. Sub-Ice Foraging Behavior of Emperor Penguins. Journal of Experimental Biology, 203: 3275? 3278. Ponganis, P. J., R. P. van Dam, D. H. Levenson, T. Knower, and K. V. Ponganis. 2004. Deep Dives and Aortic Temperatures of Emperor Penguins: New Directions for Biologging at the Isolated Dive Hole. Memoirs of National Institute of Polar Research Special Issue No. 58, Biologging Science, pp. 155? 161. Siniff, D. B., D. P. Demaster, R. J. Hofman, and L. L. Eberhart. 1977. An Analysis of the Dynamics of a Weddell Seal Population. Ecological Monographs, 47(3): 319? 335. Testa, J. 1994. Over-Winter Movements and Diving Behaviour of Fe- male Weddell Seals (Leptonychotes weddellii). Canadian Journal of Zoology, 72: 1700? 1710. Wienecke, B., G. Robertson, R. Kirkwood, and K. Lawton. 2007. Ex- treme Dives by Free-Ranging Emperor Penguins. Polar Biology, 30: 133? 142. 19_Kooyman_pg265-270_Poles.indd 27019_Kooyman_pg265-270_Poles.indd 270 11/17/08 9:30:04 AM11/17/08 9:30:04 AM ABSTRACT. We examined long-term variations in kelp growth in coincidence with recent (2004? 2006) measurements of underwater photosynthetically active radiation (PAR), light attenuation coeffi cients, chlorophyll concentrations, and total suspended solids (TSS) to determine the impact of sediment resuspension on the productivity of an isolated kelp bed community on the Alaskan Beaufort Sea coast. Attenuation coeffi cients exhibited distinct geographical patterns and interannual variations between 2004 and 2006 that were correlated with temporal and geographical patterns in TSS (range 3.5? 23.8 mg L H110021 ). The low chlorophyll levels (H110213.0 H9262g L H110021 ) in all three years were unlikely to have contributed signifi cantly to periods of low summer water transparency. Blade elongation rates in the arctic kelp, Laminaria solidungula, are excellent integrators of water transparency since their annual growth is completely dependent on PAR received during the summer open-water period. We noted that blade growth at all sites steadily increased between 2004 and 2006, refl ective of increased underwater PAR in each succes- sive year. Mean blade growth at all sites was clearly lowest in 2003 (H110218 cm) compared to 2006 (18? 47 cm). We attribute the low growth in 2003 to reported intense storm activ- ity that likely produced extremely turbid water conditions that resulted in low levels of ambient light. Examination of a 30-year record of annual growth at two sites revealed other periods of low annual growth that were likely related to summers characterized by exceptional strong storm activity. Although kelp growth is expected to be higher at shal- lower sites, the reverse occurs, since sediment re-suspension is greatest at shallower water depths. The exceptionally low growth of kelp in 2003 indicates that these plants are liv- ing near their physiological light limits, but represent excellent indicators of interannual changes in water transparency that result from variations in local climatology. INTRODUCTION Research studies conducted over the past two decades have clearly docu- mented that kelp biomass, growth, and productivity in the Alaskan Beaufort Sea are strongly regulated by light availability (photosynthetically active radia- tion, PAR). Results from a variety of experimental studies, including the linear growth response of kelp plants to natural changes in the underwater light fi eld (Dunton, 1984; 1990; Dunton and Schell, 1986;), carbon radioisotope tracer ex- periments (Dunton and Jodwalis, 1988), and laboratory and fi eld physiological work ( Henley and Dunton, 1995; 1997) have been used successfully to develop models of kelp productivity in relation to PAR. Yet, until recently, the relationship Kenneth H. Dunton and Susan V. Schonberg, University of Texas Marine Science Institute, 750 Channel View Dr., Port Aransas, TX 78373, USA. Dale W. Funk, LGL Alaska Research Associates, Inc., 1101 East 76th Avenue, Suite B, Anchorage, AK 99516, USA. Corresponding author: K. Dun- ton (ken.dunton@mail.utexas.edu). Accepted 28 May 2008. Interannual and Spatial Variability in Light Attenuation: Evidence from Three Decades of Growth in the Arctic Kelp, Laminaria solidungula Kenneth H. Dunton, Susan V. Schonberg, and Dale W. Funk 20_Dunton_pg271-284_Poles.indd 27120_Dunton_pg271-284_Poles.indd 271 11/17/08 9:23:09 AM11/17/08 9:23:09 AM 272 SMITHSONIAN AT THE POLES / DUNTON, SCHONBERG, AND FUNK between water turbidity? as measured by total suspended solids (TSS) or optical instruments? and benthic algal pro- duction was unknown. Aumack et al. (2007) were the fi rst to establish the quantitative link between water column turbidity, PAR and kelp production through a model that uses TSS data to predict estimates of kelp productivity in an area known as the Stefansson Sound Boulder Patch on the central Alaskan Beaufort Sea coast. This information is essential for evaluating how changes in water transpar- ency are related to higher suspended sediment concentra- tions from anthropogenic activities near the Boulder Patch, coastal erosion, and increased freshwater infl ow (McClel- land et al., 2006). The quantitative measurements of TSS collected by Aumack et al. (2007) in summers 2001 and 2002 were a critical fi rst step in the establishment of an accurate basin-wide production model for the Stefansson Sound Boulder Patch. The productivity of Laminaria solidungula in subtidal coastal ecosystems is an important factor that regulates benthic biodiversity and ultimately, the intensity of biolog- ical interactions such as competition, facilitation, preda- tion, recruitment, and system productivity (Petraitis et al., 1989; Worm et al., 1999; Mittelbach et al., 2001; Paine 2002). On a larger scale, biodiversity measurements can serve as an indicator of the balance between speciation and extinction (McKinney, 1998a; 1998b; Rosenzweig, 2001). The interesting biogeographic affi nities of organ- isms in the Boulder Patch led Dunton (1992) to refer to the area as an ?arctic benthic paradox,? based on the Atlan- tic origin of many of the benthic algae (e.g. the red algae Odonthalia dentata, Phycodrys rubens, Rhodomela con- fervoides) in contrast to the Pacifi c orientation of many of the invertebrates (most polychaetes and gastropods). This unique character of the biological assemblage, combined with the Boulder Patch?s isolated location (Dunton et al., 1982), suggests the potential of the area as a biogeographic stepping-stone. Thus, the Boulder Patch likely has large biological and ecological roles outside Stefansson Sound. The overarching objective of our study was to use syn- optic and long-term measurements of PAR, light attenua- tion coeffi cients, total suspended solids (TSS; mg L H110021 ), and indices of kelp biomass to determine the impact of sediment resuspension on kelp productivity and ecosystem status in the Stefansson Sound Boulder Patch. Between 2004 and 2006, we initiated studies to monitor water quality, light, kelp growth, and the associated invertebrate community in the Boulder Patch. This research program was designed to address ecosystem change as related to anthropogenic ac- tivities from oil and gas development. Our initial effort was focused on establishing a quantitative relationship between total suspended solids (TSS) and benthic kelp productiv- ity (see Aumack, 2003; Aumack et al., 2007). Our current objectives included (1) defi ning the spatial variability in annual productivity and biomass of kelp, (2) monitoring incident and in situ ambient light (as PAR) and TSS, and (3) using historical datasets of kelp growth to establish a long-term record of kelp productivity. MATERIALS AND METHODS Our overall sampling strategy during summers 2004, 2005, and 2006 incorporated: (1) semi-synoptic maps of TSS and light attenuation parameters generated through sampling at 30 randomly-selected points in a 300 km 2 area that included the Boulder Patch and the region south of Narwhal Island to Point Brower on the Sagavanirktok Delta (Figure 1; 70?23H11032N; 147?50H11032W); (2) long-term variations in underwater PAR at three fi xed sites and incident PAR at one coastal site during the summer open-water period; and (3) kelp growth at several monitoring stations established during the 1984? 1991 Boulder Patch Monitoring Program (LGL Ecological Research Associates and Dunton, 1992). A majority of our study sites were located within the Stefans- son Sound Boulder Patch, which is characterized by non- contiguous patches of H1102210% rock cover. These patches are depicted by gray contour lines in Figures 1?5. SYNOPTIC SAMPLING In order to describe the spatial extent and patterns of TSS, light attenuation, chlorophyll, nutrients, and physiochemical properties across Stefansson Sound, we sampled 30 sites across the monitoring area (Figure 1), which ranges in depth from 3 to 7 m. The location for each site was chosen by laying a probability-based grid over the area and randomly choosing a location within each grid cell. This method allowed sampling locations to be spaced quasi-evenly across the landscape while still maintaining assumptions required for a random sample (i.e., all locations have an equal chance of being sampled). All 30 sites were visited on three separate occasions during summers 2004, 2005, and 2006 using a high-speed vessel (R/V Proteus). We measured TSS, incident PAR, inorganic nutrients (ammonia, phosphate, silicate, nitrogen), water column chlorophyll, and physiochemical parameters (tem- perature, salinity, dissolved oxygen, and pH) during each synoptic sampling effort. 20_Dunton_pg271-284_Poles.indd 27220_Dunton_pg271-284_Poles.indd 272 11/17/08 9:23:10 AM11/17/08 9:23:10 AM INTERANNUAL VARIABILITY IN LIGHT ATTENUATION 273 Replicate water samples were collected at 2 and 4 m depths using a van Dorn bottle. All samples were placed in pre-labeled plastic bottles, with sampling point geographic coordinates (Lat/Long) recorded using a handheld Garmin Global Positioning System, GPSMap 76S (Garmin Inter- national Inc., Olathe, Kansas, USA). In situ physiochemi- cal measurements were made from the vessel. All other samples were stored in a dark cooler and transported to a laboratory on Endicott Island for processing. Light Attenuation Simultaneous surface and underwater measurements of PAR data were collected using LI-190SA and LI-192SA cosine sensors, respectively, connected to a LI-1000 data- logger (LI-COR Inc., Lincoln, Nebraska, USA). The LI- 190SA sensor was placed at a 4 m height on the vessel mast. Coincident underwater measurements with the LI-192SA sensor were made using a lowering frame deployed at 2 and 4 m depths. Care was taken to avoid interference from shading of the sensor by the vessel. The Brouger-Lambert Law describes light attenuation with water depth: kII z oz = ln( / ) (a) where I o is incident (surface) light intensity, I z is light in- tensity at depth z, and k is the light attenuation coeffi cient (m H110021 ). TSS A known volume of water from each sample was fi l- tered through pre-weighed, pre-combusted glass fi ber fi lters (Pall Corporation, Ann Arbor, Michigan, USA). Following a distilled water rinse fi lters were oven-dried to constant weight at 60?C. The net weight of particles collected in each sample was calculated by subtracting the fi lter?s initial weight from the total weight following fi ltration. Chlorophyll For chlorophyll measurement, 100 ml of water from each replicate sample was fi ltered through a 0.45 H9262m cellulose nitrate membrane fi lter (Whatman, Maidstone, England) in darkness. After fi ltration, the fi lters and residue were placed in pre-labeled opaque vials and frozen. The frozen fi lters were transported to The University of Texas Marine Science Institute (UTMSI) in Port Aransas, Texas, for chlorophyll analysis. At UTMSI, fi lters were removed from the vials and placed in pre-labeled test tubes containing 5 ml of metha- nol for overnight extraction ( Parsons et al., 1984:3? 28). Chlorophyll a concentration, in H9262g L H110021 , was determined using a Turner Designs 10-AU fl uorometer (Turner Design, Sunnyvale, California, USA). Non- acidifi cation techniques are used to account for the presence of chlorophyll b and phaeopigments ( Welschmeyer, 1994). Nutrients Water samples were frozen and transferred to UTMSI for nutrient analysis. Nutrient concentrations for NH 4 H11001 , PO 4 3H11002 , SiO 4 , and NO 2 H11002 H11001 NO 3 H11002 were determined by continuous fl ow injection analysis using colorimetric tech- niques on a Lachat QuikChem 8000 (Zellweger Analytics Inc., Milwaukee, Wisconsin, USA) with a minimum detec- tion level of 0.03 H9262M. FIGURE 1. The project study area showing 30 synoptic collection sites used in summers 2004, 2005, and 2006. SDI: Satellite Drilling Island. Gray contour lines show H1102210% rock cover. 20_Dunton_pg271-284_Poles.indd 27320_Dunton_pg271-284_Poles.indd 273 11/17/08 9:23:10 AM11/17/08 9:23:10 AM 274 SMITHSONIAN AT THE POLES / DUNTON, SCHONBERG, AND FUNK Physiochemical Parameters Temperature ( o C), salinity (?), dissolved oxygen (% and mg L H110021 ) pH, and water depth (m), were measured using a YSI Data Sonde (YSI Inc., Yellow Springs, Ohio, USA). PERMANENT SITES Underwater Irradiance In addition to the synoptic sampling, we established eight permanent sites for collection of long-term data. Continuous underwater PAR measurements were col- lected at three sampling sites (DS-11, E-1, and W-1) in the Boulder Patch study area (Figure 2) and terrestrial PAR measurements at one coastal location (Endicott Is- land). These sites have been the focus of previous long- term monitoring efforts; measurements of PAR and kelp growth are reported in published literature (Dun- ton, 1990). Site DS-11, established as a reference site, has been a primary research site for the Boulder Patch since 1978. This site lies well outside the area most likely impacted by sediment plumes originating from the pro- posed Liberty Project, including construction of a buried pipeline and Stockpile Zone 1 (Ban et al., 1999). All three Boulder Patch sites are located on seabed characterized by H1102225% rock cover. Sites were chosen based on either their southern-most location in the Boulder Patch (W-1, E-1), existence of historical PAR data (W-1, E-1, and DS- 11), and their likelihood of being impacted by oil and gas development through dredging activities associated with pipeline or island construction. Underwater data were collected using LI-193SA spher- ical quantum sensor (for scalar measurements) connected to a LI-1000 datalogger (LI-COR Inc., Lincoln, Nebraska, USA) at each site. Sensors were mounted on PVC poles and positioned just above the kelp canopy to prevent fouling or shading by kelp fronds. Instantaneous PAR measure- ments were taken at 1 min intervals and integrated over 1 h periods. Coincident surface PAR measurements were taken with LI-190SA terrestrial cosine sensor connected to LI-1000 datalogger located on Endicott Island. Kelp Elongation At each of the nine dive sites (depth range 5? 7 m) within the Boulder Patch (DS-11, Brower-1, E-1, E-2, E-3, L-1, L-2, W-1, and W-3), SCUBA divers collected 15? 30 individual specimens of Laminaria solidungula attached to large cobbles and boulders in summers 2004, 2005, and 2006. Samples were placed in pre-labeled black bags, transported to Endicott, and processed. Blade segments from every specimen, which corresponded to one year?s growth (Dunton, 1985), were measured and recorded to produce a recent (3? 4 yr) growth record of linear blade expansion at each site. Blades measured in summer refl ect growth during both the present and previous calendar year since more than 90% of a kelp?s frond expansion oc- curs between November and June under nearly complete darkness (Dunton and Schell, 1986). Linear growth in L. solidungula from the Boulder Patch is heavily dependent on photosynthetic carbon reserves that accumulate during the previous summer in proportion to the underwater light environment. A growth year (GWYR) is dictated by the formation of a new blade segment, which begins in mid- November every year and is defi ned by the summer that precedes new blade formation (e.g., basal blade growth measured in summer 2007 depicts GWYR 2006). Kelp Biomass Frond lengths of Laminaria solidungula plants were measured at W-3, E-1, E-3, and DS-11 along four 25 m FIGURE 2. The project study area. The indicated sites are historical Boulder Patch stations that have been visited repeatedly since 1984. During summers 2004, 2005, and 2006, long-term light was mea- sured at sites DS-11, E-1, and W-1; kelp blade length data were also collected at these sites. 20_Dunton_pg271-284_Poles.indd 27420_Dunton_pg271-284_Poles.indd 274 11/17/08 9:23:33 AM11/17/08 9:23:33 AM INTERANNUAL VARIABILITY IN LIGHT ATTENUATION 275 transects. Transects radiated from a central point at random chosen directions at 280?, 80?, 260?, and 110?Magnetic. Statistics and GIS TSS concentrations, chlorophyll a concentrations, and the attenuation coeffi cient (k) were matched with their respective geographic coordinates and plotted using GIS software ArcMap 9.2 (ERSI, Redlands, California). Data were interpolated across a polygon of Stefansson Sound, including the Boulder Patch, using Geospatial Analyst ex- tension and Kriging function in ArcMap following Aumack et al. (2007). Data were analyzed using standard paramet- ric models. Spatial and interannual signifi cance among k, TSS, and chlorophyll measurements were determined using a paired t-test to examine signifi cant differences (p H11021 0.05) among treatment variables using Microsoft Excel. Signifi - cant differences in PAR among years and sites was tested using a two-way analysis of variance (ANOVA) using time as a block with a general linear models procedure (SAS In- stitute Inc, 1985) following Dunton (1990). RESULTS SYNOPTIC SAMPLING Light attenuation (k) was derived from coincident in situ measurements of surface and underwater PAR at 2 and 4 m depths collected at 30 stations on three differ- ent occasions each summer. Attenuation was consistently elevated in coastal zones with highest k values observed near Endicott Island and SDI (Satellite Drilling Island) indicating more turbid water closer to shore (Figure 3). Lower k values were recorded offshore along the eastern and northeastern sides of Stefansson Sound. In summer 2004, k ranged from 0.43? 1.34 m H110021 (mean 0.73 H11006 0.14) throughout Stefansson Sound. In 2005 k ranged from 0.47? 1.32 m H110021 (mean 0.69 H11006 0.03) and in 2006, k was 0.54? 1.08 m H110021 (mean 0.72 H11006 0.01). The majority of the Boulder Patch, including areas with dense kelp popula- tions (H1102225% rock cover), were found predominantly in offshore waters where attenuation measurements were consistently less than 1.0 m H110021 . The TSS concentrations were dramatically lower in summers 2004 and 2006 compared with 2005, yet the same general trends were observed (Figure 4). Since a paired t-test indicated that the TSS values measured at 2 and 4 m depths were not signifi cantly different in either year (2004 p H11005 0.065; 2005 p H11005 0.156) the means of the two depths are displayed. In 2004, the highest concentrations (7.6? 8.3 mg L H110021 ) were found near Endicott Island and SDI and in a turbid area just north of Narwhal Island (5.7? 6.1 mg L H110021 ). The TSS ranged from 3.8? 7.6 mg L H110021 outside the Boulder Patch with a mean of 5.0 mg L H110021 . Inside the Boulder Patch the data ranged from 4.0 to 8.3 mg L H110021 (mean 5.0 mg L H110021 ); the overall site average was 5.0 H11006 1.8 mg L H110021 . The TSS measurements were much higher and var- ied greatly throughout Stefansson Sound during summer FIGURE 3. Combined mean attenuation coeffi cient (k) values calculated from measurements collected at 2 m and 4 m water depths in summers 2004, 2005, and 2006. 0.41 - 0.50 k (m-1) 0.51 - 0.60 0.61 - 0.70 0.71 - 0.80 0.81 - 0.90 0.91 - 1.00 1.01 - 1.10 1.11 - 1.20 1.21 - 1.30 1.31 - 1.40 20_Dunton_pg271-284_Poles.indd 27520_Dunton_pg271-284_Poles.indd 275 11/17/08 9:23:56 AM11/17/08 9:23:56 AM 276 SMITHSONIAN AT THE POLES / DUNTON, SCHONBERG, AND FUNK 2005 (7.5? 23.8 mg L H110021 ; mean 11.1 H11006 1.1 mg L H110021 ). The highest values (17.6? 23.8 mg L H110021 ) were located nearshore, adjacent to Endicott Island, SDI, and Point Brower. Out- side the Boulder Patch, TSS ranged from 7.5 to 23.8 mg L H110021 (mean 11.2 mg L H110021 ). Inside the Boulder Patch, values ranged from 9.0 to 17.6 mg L H110021 (mean 11.0 mg L H110021 ). In 2006, TSS concentrations were similar to those mea- sured in 2004 (range of 3.5? 6.9 mg L H110021 ; mean of 4.7 H11006 0.2 mg L H110021 ). The highest values were again adjacent to Endicott Island, SDI and Point Brower. Outside the Boulder Patch, TSS ranged from 3.6 to 6.9 mg L H110021 (mean 4.6 H11006 0.2 mg L H110021 ). The TSS values ranged from 3.5 to 5.9 mg L H110021 inside the Boulder Patch with a mean of 4.6 H11006 0.2 mg L H110021 . Chlorophyll a measurements from 2 and 4 m depths were relatively low but signifi cantly different from each other in all years (p H11021 0.05). In all three years, 4 m chlo- rophyll values were higher than the 2 m measurements (Figure 5). The 2005 chlorophyll means were the highest followed by 2004 means, with the lowest values occurring in 2006. In 2004, chlorophyll measurements ranged from 0.11 to 2.63 H9262g L H110021 (mean 0.39 H11006 0.2 H9262g L H110021 ). In summer 2005, values ranged from 0.11 to 3.54 H9262g L H110021 (mean 0.76 H11006 0.08 H9262g L H110021 ) compared to 0.11? 0.41 H9262g L H110021 (mean 0.18 H11006 0.01 H9262g L H110021 ) in 2006. Ammonium concentrations (Table 1) were signifi - cantly different among samples collected at 2 and 4 m in all years (p H11021 0.05); all values were low (0.12 H11006 0.06 H9262M at 2 m and 0.17 H11006 0.07 H9262M at 4 m in 2004; 0.40 H11006 0.04 H9262M at 2 m and 0.65 H11006 0.04 H9262M at 4 m in 2005 and 0.25 H11006 0.05 H9262M at 2 m and 0.12 H11006 0.02 H9262M at 4 m). Ammonium ranged from 0.0? 0.52 H9262M in 2006. Highest concentrations were noted at sites adjacent to barrier is- lands (Sites 13 and 15); lowest values were noted offshore (0.0? 0.02 H9262M). In general, phosphate concentrations were low. Mean values in 2004 were 0.24 H11006 0.03 H9262M at 2 m and 0.19 H11006 0.05 H9262M at 4 m. Phosphate values ranged from 0.11? 0.39 H9262M with the highest concentrations collected at sites ad- jacent to Endicott. Several other random sites displayed higher values at either 2 or 4 m. The lowest values were ob- served at sites seaward of Narwhal Island (0.0? 0.02) H9262M. In 2005, phosphate measurements were lower in 2 m sam- ples (mean 0.29 H11006 0.01 H9262M) versus the 4 m samples (mean 0.35 H11006 0.01 H9262M). The same pattern held for the 2006 (2 m mean 0.17 H11006 0.01 H9262M; 4 m mean 0.20 H11006 0.01 H9262M). Silicate values collected from 2 and 4 m in 2004 were not signifi cantly different (p H11005 0.51), ranging from 0.07? 4.90 H9262M; mean 1.84 H11006 0.26 H9262M (Table 1). Silicate was quite low at 2 and 4 m in 2004 compared to 2005 (2 m mean 5.64 H11006 0.19 H9262M; 4 m mean 5.19 H11006 0.16 H9262M) and 2006 (2 m mean 7.05 H11006 0.14 H9262M; 4 m mean 6.89 H11006 0.16 H9262M). NO 2 H11002 H11001 NO 3 H11002 measurements throughout Stefansson Sound were also generally low, with 2004 station means ranging from 0.0? 0.29 H9262M at 2 m; 0.12 H11006 0.21 H9262M at 4 m (Table 1). The 2005 station means ranged from 0.0? 0.61 H9262M at 2 m; 0.03? 1.98 H9262M at 4 m, and 2006 means were 0.03? 0.33 H9262M at 2 m; 0.02? 0.44 H9262M at 4 m. In all three 3.01 - 4.00 TSS (mg/L) 4.01 - 5.00 5.01 - 6.00 6.01 - 7.00 7.01 - 8.00 8.01 - 9.00 9.01 - 11.00 11.01 - 13.00 13.01 - 15.00 15.01 - 17.00 17.01 - 19.00 19.01 - 21.00 21.01 - 23.00 FIGURE 4. Combined mean total suspended solids (TSS) from samples collected at 2 m and 4 m water depths in 2004, 2005, and 2006. 20_Dunton_pg271-284_Poles.indd 27620_Dunton_pg271-284_Poles.indd 276 11/17/08 9:24:33 AM11/17/08 9:24:33 AM INTERANNUAL VARIABILITY IN LIGHT ATTENUATION 277 0.00 - 0.25 Chl (H9262g/L) 0.26 - 0.50 0.51 - 0.75 0.76 - 1.00 1.01 - 1.25 1.26 - 1.50 1.51 - 1.75 1.76 - 2.00 2.01 - 2.25 2.26 - 2.50 2.51 - 2.75 FIGURE 5. Chlorophyll a (chl) values measured in 2004, 2005, and 2006. Samples were collected at (top) 2 m and (bottom) 4 m water depth. TABLE 1. Measurements of ammonium, phosphate, silicate, and nitrate H11001 nitrite at 30 sites measured annually in July and August 2004, 2005, and 2006. Samples were collected at 2 and 4 m water column depths. Values are x H11006 SE. Nitrate H11001 Nitrate H11545 Phosphate Phosphate Silicate Silicate Nitrite Nitrite YEAR Ammonium Ammonium (PO 4 3H11546 ) (PO 4 3H11546 ) (SiO 4 ) (SiO 4 ) (NO 2H11546 H11545 (NO 2H11545 H11545 H9262M (NH 4 H11545 ) 2 m (NH 4 H11545 ) 4 m 2 m 4 m 2 m 4 m NO 3H11546 ) 2 m NO 3H11546 ) 4 m 2004 0.12 H11006 0.06 0.17 H11006 0.07 0.24 H11006 0.03 0.19 H11006 0.05 1.76 H11006 0.18 1.84 H11006 0.26 0.14 H11006 0.10 0.15 H11006 0.01 2005 0.40 H11006 0.04 0.65 H11006 0.05 0.29 H11006 0.01 0.35 H11006 0.01 5.64 H11006 0.19 5.19 H11006 0.16 0.21 H11006 0.04 0.29 H11006 0.08 2006 0.25 H11006 0.05 0.12 H11006 0.02 0.17 H11006 0.01 0.20 H11006 0.01 7.05 H11006 0.14 6.89 H11006 0.16 0.07 H11006 0.01 0.10 H11006 0.02 20_Dunton_pg271-284_Poles.indd 27720_Dunton_pg271-284_Poles.indd 277 11/17/08 9:25:15 AM11/17/08 9:25:15 AM 278 SMITHSONIAN AT THE POLES / DUNTON, SCHONBERG, AND FUNK sampling years, the 4 m nitrate concentrations were slightly higher than the 2 m but were not signifi cantly different. Mean sea surface temperature (2 m and 4 m) increased throughout the Boulder Patch each year between 2004 and 2006 (Table 2). Summer 2004 was characterized by fre- quent storm activity, which was refl ected in depressed sur- face water temperatures that were negative at some sites. The 2006 mean 2 m temperature (4.6 H11006 0.2?C) was more than double the value measured in 2004 (2.1 H11006 0.6?C). The 4 m mean temperature increased more than fourfold between 2004 and 2006 (0.9 H11006 0.7; 4.2 H11006 0.2). Salinity measurements were homogeneous across the Boulder Patch and means were consistent between sum- mers 2004 and 2005 at both 2 m (23.8 H11006 1.0?; 23.8 H11006 1.7 ?) and 4 m (26.8 H11006 1.2 ?; 26.6H11006 1.4 ?) depths, but values dropped precipitously in 2006 (2 m 16.9 H11006 0.35?; 4 m 20.7H11006 0.5?; Table 2). In 2004, the salinity range at 2 m was 20.4? 27.4?; the 4 m 2004 range was 23.5 to 31.7?. In summer 2005, measurements at 2 m ranged from 17.7 to 26.21? and at 4 m salinity varied from 24.9 to 30.8?. During 2006, the 2 m salinity low was mea- sured at 11.3 ? and the high was 23.5?; the 4 m low was 12.3 ? and high 30.0?. At 2 m, waters were slightly fresher than at 4 m during all three years. We only report pH data from 2004 and 2006 since the probe malfunctioned during the 2005 fi eld season (Table 2). In 2004, pH measurement means were remarkably constant at 8.2 H11006 0.04 at both 2 and 4 m. The measure- ments in 2006 were also very consistent (7.9 H11006 0.01 ?) throughout the sampling area and between the 2 and 4 m depths, but were more acidic than the 2004 values. Mea- surements of dissolved oxygen revealed values at or near saturation in all years (Table 2). The range in values refl ect differences in water temperature and wind induced turbu- lence of surface waters. PERMANENT STATIONS Blade elongation in Laminaria solidungula displayed large spatial and temporal variability as refl ected in mea- surements from nine sites (Table 3). Mean site blade growth was lower at every site in GWYR 2003 compared to GWYRs 2004, 2005, and 2006, refl ecting the excep- tionally poor weather conditions in summer 2003 that produced extremely low levels of ambient PAR. Kelp col- lected in 2004 (GWYR 2003) had annual average growth rates ranging between 4.4 and 10.4 cm. In contrast, blade elongation ranged from 10 to 25.5 cm in GWYR 2004 and from 16.7 to 47.3 cm in GWYR in 2005 and 2006, comparable to previous studies (Dunton, 1990; Martin and Gallaway, 1994). Specimens from DS-11 showed the greatest annual growth of all sites in most years. In conjunction with blade elongation rates, we col- lected PAR measurements during summers 2004, 2005, and 2006. Surface irradiance followed a typical cyclical pattern, peaking between 1200? 1400 H9262mol photons m H110022 s H110021 in the afternoon before declining to nearly undetect- able levels after midnight (Figure 6). Highest levels of un- derwater PAR were normally recorded by scalar sensors in the early afternoon (1400 hrs), which also corresponded to the period of maximum incident PAR. In 2004, from 31 July to 6 August, underwater irradiance dropped to near zero at all three sites (DS-11, E-1, and W-1). These low values were coincident with a series of intense storms that generated winds in excess of 10 m s H110021 from the southwest and southeast. Extremely low underwater PAR concentra- tions continued through 9 August followed by four days of slightly higher values, at which point the dataloggers were removed. Prior to the storm, underwater scalar ir- radiance at DS-11 ranged from 180 to 200 H9262mol m H110022 s H110021 compared to less than 20 H9262mol m H110022 s H110021 during the storm, TABLE 2. Average temperature, salinity, dissolved oxygen, and pH measurements for 30 sites measured annually in July and August 2004, 2005, and 2006. Samples were collected at 2 and 4 m water column depths. Values are means H11006 SE. ND indicates no data. Temp Temp Salinity Salinity Dissolved Dissolved pH pH (?C) (?C) (?) (?) O 2 (mg L H110021 ) O 2 (mg L H110021 ) (m H110021 ) (m H110021 ) YEAR 2 m 4 m 2 m 4 m 2 m 4 m 2 m 4 m 2004 2.11 H11006 0.55 0.88 H11006 0.75 23.80 H11006 0.99 26.81 H11006 1.16 13.18 H11006 0.35 14.22 H11006 0.38 8.19 H11006 0.05 8.22 H11006 0.04 2005 2.62 H11006 1.07 1.97 H11006 1.32 23.85 H11006 1.66 26.65 H11006 1.36 11.48 H11006 0.45 11.53 H11006 0.39 ND ND 2006 4.64 H11006 0.16 4.21 H11006 0.22 16.91 H11006 0.35 20.67 H11006 0.49 11.45 H11006 0.02 11.57 H11006 0.05 7.90 H11006 0.01 7.88 H11006 0.01 20_Dunton_pg271-284_Poles.indd 27820_Dunton_pg271-284_Poles.indd 278 11/17/08 9:26:40 AM11/17/08 9:26:40 AM INTERANNUAL VARIABILITY IN LIGHT ATTENUATION 279 TABLE 3. Average Laminaria solidungula basal blade length from nine sites in Stefansson Sound. Blade lengths were measured during summers 2004? 2007. A growth year (GWYR) is defi ned as the period beginning 15 November one year and ending 15 November the following year. Values are means H11006 SE. ND is no data. GWYR DS-11 cm E-1 cm E-2 cm E-3 cm L-1 cm L-2 cm B-1 cm W-1 cm W-3 cm 2003 7.20 H11006 0.50 4.38 H11006 0.36 7.85 H11006 0.40 5.24 H11006 0.40 6.56 H11006 0.54 6.53 H11006 0.86 7.16 H11006 0.60 7.93 H11006 0.51 10.45 H11006 0.94 2004 25.46 H11006 0.93 10.93 H11006 0.37 9.98 H11006 0.38 22.79 H11006 0.76 18.03 H11006 0.59 14.67 H11006 0.59 18.59 H11006 0.69 13.73 H11006 0.53 20.88 H11006 1.04 2005 27.65 H11006 0.85 18.97 H11006 0.53 19.10 H11006 0.76 28.54 H11006 0.93 23.77 H11006 0.91 21.82 H11006 0.84 24.76 H11006 1.02 18.49 H11006 0.71 24.94 H11006 1.05 2006 47.32 H11006 1.64 16.71 H11006 0.71 18.04 H11006 0.96 25.84 H11006 1.07 18.85 H11006 0.75 36.77 H11006 1.72 20.70 H11006 1.00 ND ND FIGURE 6. Continuous measurements of surface and underwater PAR in Stefansson Sound in summers 2004, 2005, and 2006. Water depths ranged from 5 m (E-1) to 6.5 m (DS-11 and W-1). Missing surface PAR data in 2004 and 2005 were obtained from an irradiance sensor main- tained 5 km distant at SDI by Veltkamp and Wilcox (2007). 20_Dunton_pg271-284_Poles.indd 27920_Dunton_pg271-284_Poles.indd 279 11/17/08 9:26:41 AM11/17/08 9:26:41 AM 280 SMITHSONIAN AT THE POLES / DUNTON, SCHONBERG, AND FUNK before rebounding to about 100 H9262mol m H110022 s H110021 . In 2005, underwater PAR was lowest at E-1 and W-1 from 26? 31 July in response to a storm event that generated winds in excess of 9 m s H110021 from the east-northeast but which had little effect on underwater PAR at DS-11. Overall, water transparency, as refl ected by consis- tently low k values (generally H110211.0 m H110021 ) and high light transmission (H1102255% m H110021 ) at all three sites, was highest in 2006 as refl ected by the absence of signifi cant storm events during the study period (Figure 7). In all three years, mean irradiance was signifi cantly (p H11021 0.05) lower at site W-1 compared to all other sites (Table 4) for the period 26 July to 10 August although the surface irradiance was high on most days. Values recorded from both surface and under- water PAR sensors are similar to irradiance measurements made in Stefansson Sound during previous studies ( Dunton, 1990). Lowest light transmission (H1102110% m H110021 ) and high- est k values (2? 3 m H110021 ) were observed at all three sites in 2004. Conditions in 2005 improved considerably, with just one peak in water turbidity occurring in late July as noted earlier. The shallower depth at E-1, compared to W-1 and DS-11 amplifi es the k values at this site for similar levels of underwater PAR recorded at all three sites. DISCUSSION The relatively uniform and low water column chloro- phyll a concentrations measured across Stefansson Sound from 2004? 2006 (Figure 5) agree well with earlier assess- ments made through the same area in 2001, but were lower than those recorded in 2002 by Aumack (2003). Chloro- phyll was consistently lower at 2 m than at 4 m depths, which may refl ect the consistently higher availability of in- organic nutrients at depth in all three years for silicate, am- monium, phosphate, and nitrate H11001 nitrite (Table 1). Water FIGURE 7. Measurements of water transparency at various sites in Stefansson Sound from 2004 to 2006. Top panel: percentage of surface irradiance (%SI). Bottom panel: diffuse attenuation coeffi cient expressed as k-values (left axis) and as % m H110021 (right axis). Underwater measure- ments were made at kelp canopy levels at E-1 (4.6 m), W-1 (5.8 m), and DS-11 (6.1 m) with a spherical quantum sensor. 20_Dunton_pg271-284_Poles.indd 28020_Dunton_pg271-284_Poles.indd 280 11/17/08 9:26:51 AM11/17/08 9:26:51 AM INTERANNUAL VARIABILITY IN LIGHT ATTENUATION 281 temperatures were about 2?C warmer in 2006 compared to 2004 and 2005, which was coincident with a 6? 7? drop in surface and bottom water salinities in 2006. Decreased sa- linities and pH in 2006 likely refl ect freshwater input from the nearby Sagavanirktok River, which produced a distinct brackish water layer to 4 m depths that was not evident in 2004 or 2005. Measurement of bottom salinity at depths exceeding 6 m at various sites (data not reported here) in- dicate that this brackish water layer seldom extended to the seafl oor, sparing benthic organisms at depths greater than 5 m exposure to widely fl uctuating salinities and tem- peratures. However, vertical gradients in temperature and/ or salinity were apparent all three years, producing a clear pycnocline. Based on in situ frond length measurements made on Laminaria solidungula plants in summers 2005 and 2006, we were able to make new calculations of kelp biomass at sites DS-11 (n H11005 226) and E-1 (n H11005 53). Areal biomass at each site was calculated using a correlation coeffi cient be- tween basal blade dry weight (gdw) and basal blade length (cm) developed for the Stefansson Sound Boulder Patch us- ing specimens collected between 1980 and 1984 (n H11005 912; Figure 8). Biomass at DS-11 (>25% rock cover) ranged from 5 to 45 gdw m H110022 (mean 23 gdw m H110022 ) compared to a range of 0.5 to 2.7 gdw m H110022 (mean 1.7 gdw m H110022 ) at site E-1 (10? 25% rock cover). Intermediate levels of biomass were recorded at sites W-3 (10.1 gdw m H110022 ) and E-3 (14.8 gdw m H110022 ), both designated as sites with H1102225% rock cover by Toimil (1980). The range in biomass at DS-11 is within the estimates reported by Dunton et al. (1982). Estimates of benthic biomass at sites in Stefansson Sound are critical for calculation of realistic basin-wide benthic production models in relation to changes in PAR. Blade elongation in Laminaria solidungula displays large spatial and temporal variability as refl ected in mea- surements from nine sites over the past decade (Figure 9). In addition, mean blade growth at two sites, DS-11 and E-1, made since 1977 and 1981, respectively, reveal some interesting long-term interannual variations (Figure 10). The two years of lowest growth (1999 and 2003) occurred relatively recently and coincide with summers character- ized by intense storm activity that likely produced ex- tremely turbid water conditions resulting in extremely TABLE 4. Mean PAR for each site for the period 26 July? 10 August (n H11005 1071 hourly measurements for each site) from 2004? 2006. Asterisks denote site means that are signifi cantly different (p H11021 0.05) within years. Average surface PAR for the same period is provided for reference. Mean PAR (H9262mol m H110022 s H110021 ) Year W-1 E-1 DS-11 Surface 2004 15.3* 23.3 28.0 314.9 2005 28.3* 45.1* 59.2* 356.0 2006 23.3* 48.1 42.0 347.9 FIGURE 8. Correlation between basal blade dry weight (g) and basal blade length (cm) in Laminaria solidungula. FIGURE 9. Variation in annual growth in Laminaria solidungula from 1996 to 2006 at sites occupied in the Stefansson Sound Boulder Patch. Measurements are based on blade lengths of plants collected between 2001 and 2006. Values are means H11006 SE. 20_Dunton_pg271-284_Poles.indd 28120_Dunton_pg271-284_Poles.indd 281 11/17/08 9:26:59 AM11/17/08 9:26:59 AM 282 SMITHSONIAN AT THE POLES / DUNTON, SCHONBERG, AND FUNK FIGURE 10. Mean annual linear growth of Laminaria solidungula from 1977 to 2006 at sites DS-11 (blue) and E-1 (green) in Stefansson Sound Boulder Patch. TABLE 5. Open-water meteorological data collected at SDI from 2001? 2006 for the period 10 July? 9 September (from Veltkamp and Wilcox, 2007). Wind Speed m s H110021 Number of Hours Average Wind Direction Average Solar Radiation Year Average Maximum H1102210 meters s H110021 ?Mag W m H110022 2001 5.10 7.18 64 157 133.7 2002 4.67 6.50 65 174 145.8 2003 5.62 7.98 158 172 123.3 2004 5.46 7.37 119 149 150.6 2005 5.57 7.47 87 125 145.8 2006 5.12 6.94 65 148 131.5 FIGURE 11. Annual mean linear growth of Laminaria solidungula as a function of the number of hours that wind speed exceeded 10 m s H110021 at SDI in July and August, from 2001 to 2006. Wind speed data from Veltcamp and Wilcox, 2007. Sites DS-11 (blue) and E-1 (green) are located in the Stefansson Sound Boulder Patch. low levels of ambient light. Wind speed data collected at SDI by Veltcamp and Wilcox (2007) from 2001 to 2006 revealed that 2003 was marked by the highest maximum wind speeds and lowest light levels in July and August when compared to all other years (Table 5). In addition, since light attenuation was lowest in offshore waters and increased with proximity to the coastline, kelp growth at site E-1 was consistently much less than at DS-11 (Table 3). Changes in local climatology clearly has an important role in regulating kelp growth as a consequence of in- creased cloud cover and sustained winds that negatively impact kelp growth (Figure 11). The exceptionally low growth of kelp in 2003 (4? 7 cm) indicates that kelp in the Boulder Patch are living close to their physiological light limits and might die if subjected to multiple years of low water transparency. Other factors that could exacerbate light limitation include increases in temperature and lower salinities. As noted above, be- tween 2004 and 2006 mean water temperature over the study area more than doubled and salinity measurements dropped signifi cantly to depths of 4 m. Since much of the Boulder Patch occurs at depths less than 6 m, kelp could be exposed to periods of lower salinities and higher tem- peratures during periods of higher than normal precipita- tion and/or freshwater infl ows. Thus, continuous monitor- ing of kelp growth in Stefansson Sound provides valuable insights into the role of local climate in affecting water transparency through processes that suspend and/or pro- mote phytoplankton (chlorophyll) production. 20_Dunton_pg271-284_Poles.indd 28220_Dunton_pg271-284_Poles.indd 282 11/17/08 9:27:13 AM11/17/08 9:27:13 AM INTERANNUAL VARIABILITY IN LIGHT ATTENUATION 283 We derived measures of light attenuation from both synoptic measurements collected at the 30 survey sites, and continuously from dataloggers deployed on the seabed. These coincident measurements of surface and underwater light exhibited distinct geographical patterns and interan- nual variations between 2004 and 2006 (Figures 3 and 7). Attenuation was consistently elevated in coastal zones, with highest values observed near Endicott Island and SDI indi- cating more turbid water closer to shore. Lower values were recorded offshore along the eastern and northeastern sides of Stefansson Sound. Attenuation coeffi cients were also highest in shallower water depths as refl ected by site E-1, compared to the lower k values at the deeper sites (W-1 and DS-11). The higher k values recorded at the permanent sites refl ect the value of continuously recording instruments; dur- ing major storms it is virtually impossible to conduct fi eld measurements, but these events are perhaps the most inter- esting and important in computing an accurate assessment of benthic production. We noted that k values were nearly three times higher at the three permanent sites than at any point during synoptic sampling at 30 sites, despite the use of a scalar sensor in the calculation of k at the permanent sites. The application of cosine measurements of PAR from the permanent sites would have resulted in still higher val- ues for k. Our data and that of Aumack et al. (2007) strongly suggest that both the spatial and interannual variations in water transparency are correlated with TSS. In general, the majority of the Boulder Patch, including areas with dense kelp populations (H1102225% rock cover), was found predominantly in clear offshore waters where attenuation measurements were consistently less than 1.0 m H110021 . Our data show that years characterized by frequent storm ac- tivity are likely to have signifi cant impacts on annual kelp growth and production. Local climatic change that results in more frequent storm events are thus likely to have a signifi cant and detrimental impact on nearshore kelp bed community production, and could lead to large scale losses of these plants and their associated diverse epilithic and epiphytic fauna. ACKNOWLEDGMENTS We thank Ted Dunton and John Dunton (University of Texas at Austin) for their mechanical expertise and pi- loting of the R/V Proteus. Special appreciation to Brenda Konar, Katrin Iken, and Nathan Stewart (all at University of Alaska Fairbanks) for their diving assistance and with kelp collection. We are grateful to Bill Streever (BP Explo- ration) for ocean access and the kindness and hospitality extended to us by the entire Endicott facility of BP Explo- ration. Constructive comments to the paper were kindly provided by Craig Aumack (University of Alabama) and Lanny Miller (Algenol Biofuels, Inc.). 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Marine Ecology Progress Series, 185: 309? 314. 20_Dunton_pg271-284_Poles.indd 28420_Dunton_pg271-284_Poles.indd 284 11/17/08 9:27:30 AM11/17/08 9:27:30 AM ABSTRACT. The life cycle of the Antarctic krill, Euphausia superba, intersects in space and time with the expansion and contraction of annual pack ice. Consequently, the circumpolar distribution of krill has often been defi ned as generally limited to an area bounded by the maximum extent of pack ice. Pack ice has both direct and indirect effects on the life cycle of krill. During the austral winter, larval krill are found in direct associa- tion with the underside of the ice and feed on the small plants and animals that constitute the sea ice microbial community, a food source relatively abundant in winter compared to food sources in the water column. Indirectly, melting pack ice in late winter or early spring stabilizes the water column and promotes growth of the preferred food of krill, which, in turn, likely provides the fuel for egg production during the summer months. Thus, the warming trend west of the Antarctic Peninsula with attendant changes in both the timing and duration of winter ice has implications for the population dynamics of krill. Given the complexity of the habitat? life cycle interaction, research on Antarctic krill involves diverse sampling tools that are dependent on the size and habitat of krill during a particular stage of their life cycle, and the nature of the study itself. In particu- lar, and pertinent to the topic of diving in polar research, the research has been greatly enhanced by diving techniques developed to allow both observation and sampling of krill in their winter pack-ice habitat. INTRODUCTION One of the reasons that Antarctic krill, Euphausia superba, has been a fo- cus of international research in the Antarctic since the Discovery days, before World War II, is that it is viewed as a key species in the Southern Ocean ecosys- tem. Various investigators have referred to Antarctic krill, Euphausia superba, as a keystone (Moline et al., 2004) or core or key (Quetin and Ross, 2003) or dominant (Ju and Harvey, 2004) species in the pelagic ecosystem of the South- ern Ocean. The rationale for the use of these terms has been based on the facts that it is among the world?s most abundant metazoan species (Nicol, 1994) and that it is important in the diets of many of the species of the upper trophic levels (Everson, 2000). However, given the suggestion that the term keystone species only refers to those species exercising an effect on ecosystem function dispropor- tionate to abundance and thus is almost always a predator (Power et al., 1996), Antarctic krill may be more accurately defi ned as a foundation species in the Langdon B. Quetin and Robin M. Ross, Univer- sity of California at Santa Barbara, Marine Science Institute-UCSB, Santa Barbara, CA 93106-6150, USA. Corresponding author: L. Quetin (Lang- don@icess.ucsb.edu). Accepted 28 May 2008. Life under Antarctic Pack Ice: A Krill Perspective Langdon B. Quetin and Robin M. Ross 21_Quetin_pg285-298_Poles.indd 28521_Quetin_pg285-298_Poles.indd 285 11/17/08 8:47:44 AM11/17/08 8:47:44 AM 286 SMITHSONIAN AT THE POLES / QUETIN AND ROSS sense of Dayton (1972). A foundation species is one that controls community dynamics and modulates ecosystem processes such as energy fl ux, and whose loss would lead to system-wide changes in the structure and function of the ecosystem (Ellison et al., 2005). Understanding that Antarctic krill may be a foundation species in many re- gions of the pelagic Southern Ocean highlights the need to elucidate the factors affecting its population dynamics and its possible response to climate change. After introduc- ing the concept of Antarctic krill as a foundation species, and the pack ice as a habitat, we focus on what has been learned about interactions between krill life history and the pack ice habitat, as well as the importance of viewing seasonal sea ice dynamics from a krill perspective. FOUNDATION SPECIES Distribution and Biomass Several characteristics of the distribution and biomass of Antarctic krill suggest it is a foundation species. First, the distribution of Antarctic krill is circumpolar. However, abundance is patchy with highest abundances in the south- east Atlantic. Most krill are found within the area south of the northern extent of annual sea ice and within the boundaries of minimal and maximal sea ice, with the ex- ception of krill around South Georgia (Marr, 1962; Laws, 1985; Siegel, 2005). This coherence led investigators to postulate a key role for seasonal pack ice in the life cycle and population dynamics of Antarctic krill. Second, Antarctic krill often dominate the zooplankton biomass in the upper 200 m of the seasonal sea ice zone (Hopkins, 1985; Hopkins and Torres, 1988; Miller and Hampton, 1989; Ward et al., 2004; Siegel, 2005). Antarctic krill biomass was recently resurveyed in Area 48, the south- west Atlantic, during the CCAMLR 2000 Survey (Hewitt et al., 2004). From these results, acoustic estimates of the circumpolar krill biomass were estimated to be 60? 155 million metric tones (Nicol et al., 2000), within the range estimated consumed by predators (Everson, 2000; Barrera- Oro, 2002). However, krill biomass in a region varies sub- stantially? seasonally with shifts in population distribution, interannually due to variation in recruitment success during its lifespan, and within a season due to local oceanographic variables (Siegel, 2005; Ross et al., 2008). Role in Ecosystem Due in part to its high biomass and in part to the fact that there are no true prey substitutes in the seasonal sea- ice zone for upper-level predators, Antarctic krill dominate energy fl ow to upper trophic levels (Barrera-Oro, 2002). In a review of the diets of Southern Ocean birds, Croxall (1984) identifi ed both (1) species highly dependent on Antarctic krill, for example, the brushtail penguins up to 98%, and (2) species whose diet was only 16? 40% krill, namely, fl ying seabirds such as albatrosses and petrels. All Antarctic seals depend somewhat on Antarctic krill, with the exception of the elephant seal (Laws, 1984). The crab- eater seal is a specialist on these euphausiids whereas the diet of leopard seals is only about 50% krill. Baleen whales (minke, blue, fi n, sei, and humpback) feed predominantly on Antarctic krill. Lastly, both fi sh and squid (Everson 2000) are known predators of krill. In particular, the me- sopelagic myctophid Electrona antarctica is an important predator of krill (Hopkins and Torres, 1988; Lancraft et al., 1989; 1991; Barrera-Oro, 2002). One aspect of the Southern Ocean ecosystem that lends support to the characterization of Antarctic krill as a foundation species is that there is little functional redundancy in prey items for the upper-trophic-level predators in the food web. Another large and some- times biomass- dominant grazer in the zooplankton is the salp, Salpa thompsoni. However, although some fi sh are known to ingest S. thompsoni, its high water con- tent and the variation in biomass by orders of magnitude during the spring/ summer season due to its characteristic alternation of generations rends it a less desirable food item. The pelagic fi sh fauna in the Southern Ocean, a logical alternate source of food, is relatively scarce and only a minor component of the epiplankton of the Ant- arctic Ocean (Morales-Nin et al., 1995; Hoddell et al., 2000), except in the high latitude regions of the cold con- tinental shelf (high Antarctic) such as the Ross Sea or the southern Weddell Sea, the habitat of the nototheniid Pleuragramma antarcticum. In deeper pelagic waters, (H11022500 m) the mesopelagic myctophid E. antarctica is the dominant fi sh (Barrera-Oro, 2002), and is available as prey to diving birds and seals when it migrates into the upper 0? 300 m during the night (Robison, 2003; Loots et al., 2007). However, in many regions of the Southern Ocean neither of these two species would be available in high enough biomass as an alternate prey if Euphausia superba disappeared. Although Antarctic krill themselves are omnivorous and do ingest both plant and animal matter (Atkinson and Sn?der, 1997; Schmidt et al., 2006), they are very impor- tant herbivorous grazers and their growth and reproduction rates appear to be tightly linked to phytoplankton concen- trations, particularly diatoms (Ross et al., 2000; Schmidt et al., 2006; Atkinson et al., 2006). This short link between 21_Quetin_pg285-298_Poles.indd 28621_Quetin_pg285-298_Poles.indd 286 11/17/08 8:47:44 AM11/17/08 8:47:44 AM LIFE UNDER ANTARCTIC PACK ICE 287 primary producers and the upper trophic levels creates a very effi cient transfer of energy to the top-level predators. The role of Antarctic krill as one of the dominant mac- rozooplanktonic grazers, particularly of the larger phyto- plankton, suggests that grazing by krill affects phytoplank- ton community composition and is a signifi cant loss term in some years (Ross et al., 1998; Garibotti et al., 2003; Daniels et al., 2006). Daniels et al. (2006), in a network analysis of the pelagic food web on the shelf west of the Antarctic Peninsula, found that in years of high primary production and high krill abundance, more than 50% of the large phytoplankton cells were ingested by Antarctic krill. In addition, its production of large fast-sinking fecal pellets (Ross et al., 1985; Fowler and Small, 1972; Cad?e et al., 1992; Gonz?lez, 1992; Turner, 2002) enhances its con- tribution to carbon sequestration (Smetacek et al., 2004). PACK ICE HABITAT FROM KRILL PERSPECTIVE Three types of pack ice can be delineated: seasonal, perennial, and marginal (Eicken, 1992). During the an- nual growth and melt cycle, the proportion of each type varies, which in turn means that the ecological habitats provided by each vary in space and time. Seasonal pack ice is a circumpolar environment that grows each fall and shrinks each spring, covering a vast area at its great- est extent. In the winter and spring, seasonal pack ice has phytoplankton/ ice algal standing stocks that are one to three orders of magnitude higher than in the water col- umn immediately below. This annual phenomenon thus provides a source of food for grazers (microheterotrophs, copepods, euphausiids) during times in the annual cycle when food resources in the water column are low. The process of formation of seasonal pack ice involves frazil ice formation scavenging particles from the water column, congelation into pancake ice, and aggregation into ice fl oes. Once the fl oes are 0.5 to 0.7 m thick, the annual ice only thickens by processes of deformation, particularly over-rafting. The seasonal pack ice, particularly the zone of highly over-rafted ice, is a favored habitat of Antarc- tic krill (Euphausia superba) in winter (Marschall, 1988; Smetacek et al., 1990; Quetin et al., 1996; Frazer et al. 1997; 2002). The perennial pack ice, in contrast, is a mix- ture of annual and primarily second year sea ice, and the water column below perennial pack ice tends to have even lower phytoplankton concentrations than below seasonal sea ice due to the increased light attenuation. In the main body of this contribution, we will illustrate how putting one?s self into the winter habitat as a diver, or taking on the krill perspective, has allowed unique in- sight into the physiological and behavioral adaptations of Antarctic krill to winter conditions. In turn, this insight allows us to further our understanding of the impact of cli- mate change. We will focus on the increased understand- ing over the last 25 years of interactions between the krill life cycle and seasonal sea ice dynamics as gained from both long-term research conducted in the summer months and from cruises conducted during winter months. Lastly, we will put these results into the context of climate warm- ing and its effect on seasonal sea ice dynamics west of the Antarctic Peninsula. BACKGROUND OBSERVATIONS LIFE HISTORY OF ANTARCTIC KRILL Characteristics of Life Cycle Antarctic krill is a relatively large (maximum length about 60 mm) and long-lived crustacean that occurs in schools, leading Hamner et al. (1983) to suggest that it has attributes more like small fi sh such as an anchovy or sar- dine than like a zooplankter. The life cycle of this euphau- siid is complex, with 11 larval stages over the fi rst 9 to 10 months, at least 1 year as a juvenile/subadult, and then 3 to 4 years as an adult. First reproduction can be as early as the third summer (Age Class 2H11001), but may be delayed if food resources are inadequate (Ross and Quetin, 2000). Ovarian development begins in the spring, fueled by food ingested during that time and not by stored reserves ( Hagen et al., 2001). The embryos sink rapidly and hatch in deep water and the nauplii swim toward the surface (Marr, 1962; Quetin and Ross, 1984; Hofmann et al., 1992). The krill in their fi rst feeding stage (Calyptopis 1) reach the surface approximately three weeks after hatching, and must fi nd suffi cient food for continued development within a few weeks or else they die (Ross and Quetin, 1989). The larvae spend their fi rst winter in the late furcilia stages; they begin to metamorphosize into juveniles and then subadults at the end of winter and throughout the spring. Critical Periods Three facets of the infl uence of seasonal pack ice on the population dynamics of Antarctic krill have emerged from the research of multiple investigators. Here we briefl y describe how two of the three critical periods interact with the seasonal cycles within the pack ice habitat (Figure 1). First, in the austral spring as the ovary begins to de- velop, investigators postulate that the female must store 21_Quetin_pg285-298_Poles.indd 28721_Quetin_pg285-298_Poles.indd 287 11/17/08 8:47:44 AM11/17/08 8:47:44 AM 288 SMITHSONIAN AT THE POLES / QUETIN AND ROSS lipid in the ?fat body? from ongoing ingestion to reach a threshold or ovarian development cannot continue (Cuzin-Roudy, 1993; Cuzin-Roudy and Labat, 1992; Ross and Quetin, 2000; Quetin and Ross, 2001), as shown for another species of euphausiid (Cuzin-Roudy, 2000). Thus, an individual female will only reproduce during a summer following a spring with adequate food sources. Each austral spring, the retreat and melting of the seasonal pack ice sets up the conditions for marginal ice-edge blooms, providing an important and timely food resource for female krill for ovarian development and eventual spawning. Second, the larva needs to feed within 10?14 days of the time of metamorphosis into the fi rst feeding stage or it passes the point-of-no-return and will not survive even if food becomes available later (Ross and Quetin, 1989). For this critical period, the effect of seasonal sea ice is indirect, through the impact of the seasonal sea ice cycle on annual primary production and the timing of blooms (Vernet et al., 2008). Lastly, although adult krill tolerate prolonged starva- tion and could survive a winter without food (Ikeda and Dixon, 1982), larval krill have a much lower starvation tolerance (for furcilia 6, about 6 wks) (Ross and Quetin, 1991; Quetin et al., 1996; Meyer and Oettl, 2005). Sea-ice microbial communities (SIMCOs) provide larval krill an alternate food source in the winter when food in the water column is at an annual low. In winter, larval krill from the under-ice habitat are in better condition than those from open water, as measured by condition factor, lipid content and in situ growth rates (Ross and Quetin, 1991), sup- porting this concept. FIGURE 1. Life cycle of Antarctic krill with three critical periods; two are directly infl uenced by seasonal sea ice dynamics: ovarian development in the austral spring and survival during the fi rst winter. 21_Quetin_pg285-298_Poles.indd 28821_Quetin_pg285-298_Poles.indd 288 11/17/08 8:47:45 AM11/17/08 8:47:45 AM LIFE UNDER ANTARCTIC PACK ICE 289 Variability in the environment, including seasonal sea ice dynamics, impacts food available to the Antarctic krill during these critical periods and is a primary fac- tor driving variability in recruitment success or year class strength. Recruitment in this species shows high interan- nual variability as illustrated by two long-term research programs in the region of the Antarctic Peninsula? Ant- arctic Marine Living Resources (AMLR), Siegel and Loeb (1995), and Palmer Long Term Ecological Research (LTER), Quetin and Ross (2003)? and by shorter series in other regions (Watkins, 1999; Siegel 2000, 2005). Within the Palmer LTER study region, recruitment has been episodic with two sequential high years followed by two to three years of low or zero recruitment (Quetin and Ross, 2003), a 5? 6 year cycle. Further north, with a longer time series, the frequency of high recruitment years has not been as repetitive (Siegel and Loeb, 1995; Siegel, 2005), although there is a rough correspondence between successful recruitment years between the two re- gions 800 km apart (Siegel et al., 2003; Ducklow et al., 2007). However, with several possible reproductive sum- mers, Antarctic krill would not need successful recruit- ment every year, and models suggest that several years of low to zero recruitment would not preclude recovery of the stock (Priddle et al., 1988). These results provide the opportunity to use correla- tions with environmental variability to formulate potential mechanisms that drive the variability (Siegel and Loeb, 1995; Quetin and Ross, 2003; Quetin and Ross, 2001). From these long-term studies, correlations have been found between both reproduction and recruitment success and various aspects of seasonal sea ice dynamics, including tim- ing, duration, and maximum extent, with the dominant parameter varying with the study region and/or latitude. Evidence from two long-term studies suggests that timing and extent of sea ice in winter and/or spring impact the re- productive cycle (Siegel and Loeb, 1995; Loeb et al., 1997; Quetin and Ross, 2001). We will use examples from the Palmer LTER 1 as illustrations of the correlations found and potential mechanisms suggested. Our fi rst example illustrates the impact of the timing of sea ice retreat in the spring on the reproductive cycle. The most important factor in estimates of population fe- cundity (numbers of larvae produced in a region during the season) is the intensity of reproduction (percentage female krill in the reproductive cycle for a season), an index that can vary by a factor of 10 interannually (Quetin and Ross, 2001). The intensity of reproduction correlated with both dynamics of sea ice in spring and with annual primary pro- duction, which are both environmental factors associated with food availability, as seasonal sea ice dynamics medi- ates the availability of food in the austral spring. Intensity of reproduction was consistently low when sea ice retreat was either early (August) or late (November), and highest when retreat occurred around the climatological mean for the region (Quetin and Ross, 2001). We emphasize here that the timing of retreat infl uences the timing of the food available for ovarian development, critical for a success- ful reproductive season. As discussed by Cuzin-Roudy (1993), Cuzin-Roudy and Labat (1992), and Quetin and Ross (2001), accumulation of stores in the ?fat body? by late spring is hypothesized to be necessary for continued ovarian development. If development does not or cannot continue because of lack of adequate food in the spring, then the intensity of reproduction is low. In the second example, we examine recruitment suc- cess and timing of sea ice advance. Our measure of recruit- ment success, the recruitment index, R 1 , is a consequence of both the numbers of larval krill entering the winter (reproductive output in the summer) and larval mortal- ity during the winter (availability of winter food sources), and thus refl ects two critical periods in the life cycle. From catches of krill from each station during the summer cruise, we calculate a recruitment index for the year, the propor- tion of one-year-old krill of the entire population one-year and older, as described in Quetin and Ross (2003). For the time series to date (Quetin and Ross, 2003; Ducklow et al., 2007) R 1 decreases exponentially with delay in sea ice advance. With advance in April, R 1 is greater than 0.4 (defi ned as a high recruitment year; Ducklow et al., 2007), but if advance is delayed until May or June, then R 1 is usually below 0.4. The suggestion is that if sea ice does not advance until late in the fall, that is, May, then recruit- ment tends to be low. However, the outliers or exceptions also provide insight into the mechanisms involved, in this instance the year classes of 1992 and 2001, as detailed in Quetin and Ross (2003). For the year class of 1992, although sea ice advanced early (March), retreat was also early (July), so SIMCOs were not available as an alternate food source in later winter, and presumably larval mortal- ity was high. For the year class of 2001, sea ice did not advance until July, yet the R 1 (0.9) indicated a very suc- cessful year class. In this year, we had observed a strong re- productive output in the summer, leading to high numbers of larval krill entering the winter, so even with presumed high mortality due to a lack of SIMCOs early in the win- ter, enough larvae survived for a strong year class. This latter point emphasizes the importance of understanding the reproductive cycle and population fecundity, as well as mortality during the fi rst year. 21_Quetin_pg285-298_Poles.indd 28921_Quetin_pg285-298_Poles.indd 289 11/17/08 8:47:56 AM11/17/08 8:47:56 AM 290 SMITHSONIAN AT THE POLES / QUETIN AND ROSS DIVING IN THE PACK ICE The above examples illustrate how seasonal sea ice dynamics is correlated with the population dynamics of Antarctic krill. What have we learned about the interac- tion between Antarctic krill and the pack ice habitat from entering the habitat itself? Historical Overview The pack ice environment is dynamic? on both sea- sonal and shorter time scales? which creates challenges for investigators. In the early 1960s, biologists began to realize that sea ice presents a variety of different modes and contains distinct communities of plants and animals (Fogg, 2005). Scuba diving with observations of both the habitat and its inhabitants has played a key role in reveal- ing the mysteries of the seasonal pack ice habitat, and scuba diving has become a key tool in investigations of the pack ice environment. Bunt (1963) used scuba diving to examine sea ice algal communities in situ and suggested ice algae could add appreciably to primary production in the Southern Ocean. He gave one of the earliest hints of the possible importance of sea ice algae as a food source for grazers. Early observations of the pack ice habitat were infrequent due to the lack of dedicated ice-capable research vessels. This limitation was relieved with the introduction of the RVIB Polarstern, commissioned in 1982 and op- erated by the Alfred Wegener Institute of Germany, and shortly thereafter, the MV Polar Duke, which began op- erations for the National Science Foundation of the USA in 1985. The advent of dedicated, ice-worthy research ves- sels led to a proliferation of studies in ice-covered waters (Ross and Quetin, 2003). Some of the earliest observations of Antarctic krill un- der the pack ice were made by U.S. Coast Guard (USCG) scuba divers during spring (November 1983) and fall (March 1986) cruises in the Weddell Sea for the Antarctic Marine Ecosystem Ice Edge Zone (AMERIEZ) program (Daly and Macaulay, 1988; 1991). Subsequently, in late winter 1985 west of the Antarctic Peninsula during the fi rst of a series of six winter cruises (WinCruise I, Quetin and Ross, 1986), divers investigating the SIMCOs associ- ated with the underside of the ice (Kottmeier and Sullivan, 1987) observed larval krill in the under-ice habitat. Quetin and Ross (1988) began research on the physiology and distribution of larval krill found on the underside of the ice with WinCruise II in 1987; recently Quetin and Ross (2007) published detailed protocols, based on their experi- ence, for diving in pack ice under various conditions that included a table of the year and month of their pack ice diving activities (Table 1). O?Brien (1987) observed both Antarctic krill and ice krill (Euphausia crystallorophias) in the under-ice habitat in austral spring of 1985. Hamner et al. (1989) found larval krill in austral fall 1986 associ- ated with newly forming sea ice. In all cases, the investiga- tors observed larval krill in higher abundance associated with the sea ice than with the water column, and observed larval krill feeding on the sea ice algae (Table 1). Investi- gators made complementary observations onboard ships both west of the Antarctic Peninsula (Guzman, 1983) and in the Weddell Sea in spring (Marschall, 1988). Gains from Diving Activities Distinct advances in our understanding of the interac- tion of Antarctic krill and the pack ice environment emerged from diving activities. Not only were larval krill observed directly feeding on sea ice algae (as detailed above), but scuba observations also documented that a clear habitat segregation existed between adult and larval krill in winter (Quetin et al., 1996), with adult krill away from the underside of the pack ice, and larval krill coupled to the underside of the pack ice. These observations led to the concept of ?risk-balancing? as put forth by Pitcher et al. (1988) for these two life stages of krill in winter; for example, the degree of association with the under-ice sur- face and its SIMCOs (food source) is a balance between the need to acquire energy and the need to avoid preda- tion. The two life stages differ in both starvation tolerance and vulnerability to predation. The smaller larvae appear to have a refuge in size (Hamner et al., 1989), as most vertebrate predators ingest primarily adults (Lowry et al., 1988; Croxall et al., 1985). Thus, the risk of predation for the adults is higher near the pack ice that is used as a plat- form for many upper-level predators. Quantitative surveys also revealed that larval krill occurred in over-rafted pack ice and not smooth fast ice, and that they were more com- monly found on the fl oors of the ?caves? formed by the over-rafting pack ice than the walls or ceilings (Frazer, 1995; Frazer et al., 1997; 2002). Not only were gains made in our understanding of the natural history and hab- itat use of Antarctic krill in winter, but also the ability to collect krill directly from the habitat has advantages over other collection methods such as towing through ice. First, the gentle collection of specimens by scuba divers with aquarium nets yields larval krill in excellent physiological conditions for experiments, for example, growth and graz- ing. Second, this method allows for immediate processing of larval krill for time-dependent indices such as pigment 21_Quetin_pg285-298_Poles.indd 29021_Quetin_pg285-298_Poles.indd 290 11/17/08 8:47:56 AM11/17/08 8:47:56 AM LIFE UNDER ANTARCTIC PACK ICE 291 content, an index of feeding on ice algae in situ (Ross et al., 2004). Entry into the under-ice habitat also meant that larval krill and their food resource (SIMCOs on the bot- tom surface of the pack ice) could be collected simultane- ously, allowing for close temporal/spatial linkages. PROCESS CRUISES? SOUTHERN OCEAN GLOBEC ICE CAMPS With the correlations that suggested mechanisms and the scuba diving protocols in place, the next step was to move beyond the correlations and explore and test mecha- nisms consistent with the observations. The evolution of pack ice diving is far from complete, however. Although long-term ice camps have been occupied in the perennial ice of the Weddell Sea on fl oes of much greater dimension than we describe below (Melnikov and Spiridonov 1996), shorter term ice camps on smaller fl oes west of the Ant- arctic Peninsula had not been attempted. On some recent cruises with both the Palmer LTER and U.S. Southern Ocean GLOBEC (Global Ocean Ecosystem Dynamics) programs, we pulled together many historical observa- tions of ways to cope with the dynamics of the pack ice environment west of the Antarctic Peninsula, and devel- oped the ability to dive from small (H1102250 m) consolidated fl oes west of the Antarctic Peninsula repeatedly for periods up to nine days (Ross et al., 2004; Quetin et al., 2007). These ice camps entailed sampling from consolidated pack ice fl oes occupied for periods of days using the research vessel to stage operations (Quetin and Ross, 2007). Div- ing on fl oes for days at a time enabled us to explore local variability in the pack ice community associated with an individual fl oe over time as the fl oe drifted within the pack ice. Scuba diving was not the only activity that occurred at these ice camps. In fact, the ability to do simultaneous sampling both from the topside and underside of the ice TABLE 1. Diving projects at Palmer Station and on cruises in the Southern Ocean, 1983? 2005. Cruises Palmer Fall Winter Spring Summer Station Year A M J J A S O N D J F M Ship 1983 AZ 1984 1985 W W O?B PD 1986 H AZ PD (H) 1987 W W PD 1988 1989 W PD 1990 K PD L 1991 K W PD L 1992 L 1993 L L W L PD L 1994 W W PD L 1995 L 1996 L 1997 L 1998 1999 L LMG L 2000 A A NBP L 2001 G G L L LMG (G) NBP (L) L 2002 G G LMG L 2003 L 2004 L 2005 AZ? AMERIEZ, W? WinCruise krill studies to Ross and Quetin, K? krill studies to Ross and Quetin, H? krill studies to Hamner, O?B? krill studies to O?Brien, L? Palmer Long-Term Ecological Research project, A? Antarctic Pack Ice Seals research project, G? Southern Ocean GLOBEC, PD? M/V Polar Duke, LMG? ARSV Laurence M Gould, NBP? RVIB Nathaniel B. Palmer. 21_Quetin_pg285-298_Poles.indd 29121_Quetin_pg285-298_Poles.indd 291 11/17/08 8:47:56 AM11/17/08 8:47:56 AM 292 SMITHSONIAN AT THE POLES / QUETIN AND ROSS fl oes made for effi cient sampling and better linkage be- tween data sets. During ice camps on two winter cruises for South- ern Ocean GLOBEC west of the Antarctic peninsula in 2001 and 2002 (methods and results described in Quetin et al., 2007), total integrated chlorophyll a in multiple ice cores (Fritsen et al., 2008) was measured on the same ice fl oes where larval krill were collected by scuba divers. The amount of chlorophyll a in the ice cores was used as a proxy for the SIMCOs available as food to the larval krill living on the underside of the pack ice. Some of the larval krill were used immediately for an index of feeding (pig- ment content) (as described in Ross et al., 2004), while others were used in instantaneous growth rate experi- ments (in situ growth rate estimates as described in Ross and Quetin, 1991; Quetin et al., 2003; Ross et al., 2004). Growth of larval krill in their winter habitat refl ects their feeding history over the past three weeks to one month, and may be an indicator of their ability to survive, that is, better growth indicates higher survivorship than lower growth. Evidence for similar linkages has been found for larval fi sh; larvae in better condition will have lower mortality rates and hence lead to stronger year classes, all else being equal (Pepin, 1991; Ottersen and Loeng, 2000; Takahashi and Watanabe, 2004). CONTRAST TWO YEARS The contrast in the results for total integrated chlo- rophyll a in ice cores, pigment content in larval krill, and the in situ growth rates of larvae for 2001 and 2002 was marked (Table 2) (fi g. 5 in Quetin et al., 2007). Gener- ally, in 2002 the ice cores contained an order of magnitude more chlorophyll a than in 2001 (Fritsen et al., 2008), leading us to suggest that more food was available to lar- val krill in 2002 than 2001. The median pigment content and in situ growth rates were also higher in larval krill in 2002 than in 2001. In both years, the distribution of pigment content was skewed to the left, but in 2002 more than 70% of the samples showed higher pigment content than those of larval krill collected from under the ice in 2001. A similar difference in distribution occurred for the in situ growth rates (Table 2). In 2002, more than 60% of the growth increments were positive, whereas in 2001 only 13% were positive. The hypothesis that high concentrations of ice algae lead to higher growth rates is supported by a comparison of the in situ growth rates in the larvae and an index of feeding for larvae collected from the same place and at the same time. For this comparison, in situ growth rates in units of percent per intermolt period (the growth incre- ment) were converted to growth in units of mm d H110021 . For larvae that molted we used the median intermolt period of 30 d found for both years (Quetin et al., 2007), and individual growth increments and total lengths to estimate growth in mm d H110021 : (total length (mm) ? % IMP H110021 )/(IMP (d)). The relationship between growth and the index of feeding for the eight in situ growth experiments from both years for which we have complimentary pigment content data is exponential, similar to a functional response curve with a maximum growth rate above a threshold feeding intensity or pigment level (Figure 2). The different symbols for the two years illustrate that data from ice camps from one year alone would not have yielded as comprehensive an illustration of the relationship between the feeding index and growth rates. Larvae with very low pigment content and negative growth rates were those from 2001, whereas larvae with a range of pigment content above 0.2 H9262g chl a g wwt H110021 and with positive growth rates were those from 2002. Thus, the combined data presents strong evidence that larval krill with a higher feeding index are growing at higher rates than those with lower feeding in- dices, and that the relationship holds at the large scale of the entire cruise (Table 2), and at the smaller scale of ice camps (Figure 2) with simultaneously collected data sets. This relationship and the difference between years in the chlorophyll a in the ice cores gives support to the infer- TABLE 2. Comparison of data from ice camps in 2001 and 2002 during two winter process cruises west of the Antarctic Peninsula near Marguerite Bay: median values of integrated chlorophyll a in ice cores, pigment content in larval krill, and in situ growth rates in larval krill. Ranges are in parentheses below median values; n H11005 number of samples. Data in graph form in Quetin et al. (2007). Data type (unit) and statistic 2001 2002 Integrated Chl a (mg m H110023 ) 1 10 Pigment Content 0.148 2.594 (median H9262g chl a gwwt H110021 ) (0.074? 0.226) (0.088? 16.615) n 29 71 In situ Growth Rate H110021.31 1.54 (% intermolt period H110021 ) (H110026.10? 5.13) (H110023.23? 11.69) n 114 132 21_Quetin_pg285-298_Poles.indd 29221_Quetin_pg285-298_Poles.indd 292 11/17/08 8:47:57 AM11/17/08 8:47:57 AM LIFE UNDER ANTARCTIC PACK ICE 293 ence that larval krill have higher growth rates during years when there is more ice algae in the pack ice. INTERACTION OF KRILL POPULATION DYNAMICS AND SEA ICE DYNAMICS CONCEPTS DEVELOPED One of the major concepts developed from these re- sults and from the results of a diagnostic algal growth and ice dynamics model (Fritsen et al., 1998) is that not all pack ice has the same value as habitat for larval krill. Not only have we learned that larval krill appear to prefer over-rafted pack ice in preference to fast ice or un-rafted pack ice, but we have learned that there is signifi cant vari- ability in the quality of the habitat that the over-rafted pack ice habitat affords larval krill. What causes these differences in habitat quality for larval krill? The hypoth- esis is that the timing of ice formation in the austral fall impacts two aspects of production in SIMCOs and thus food available: (1) the amount of material in the water column to be scavenged and incorporated into the form- ing ice, or the base standing stock, and (2) the amount of photosynthetically available radiation (PAR) for in situ growth of the ice algae to take place integrated over the time between ice formation and mid-winter darkness. In general the rate of accumulation of SIMCO biomass slows during the transition from fall to winter as the daily PAR decreases (Hoshiai, 1985; Fritsen and Sullivan, 1997; Mel- nikov, 1998), following the decrease in day length. Simulations predict that in winters when ice forms early, chlorophyll a will be higher in the pack ice due to both factors. Even a 10-day delay may cause an effect (C. H. Fritsen, unpublished data). Timing of ice formation is critical since the earlier ice forms, the higher the probabil- ity of incorporation of high abundances of phytoplankton from fall blooms into the ice lattice, and earlier ice forma- tion also means more total light available for the ice algae to grow before mid-winter when light levels are too low for net primary production at most latitudes. The varia- tion in PAR is signifi cant, as illustrated by the decrease by more than 50% in the day length from March to April to May: 12.8 h to 9.2 h to 5.6 h at 66?S (Figure 3) and 12.8 h to 4.4 h at 68?S (Quetin et al. 2007). For our two-year comparison, the sea ice advance was a month earlier in 2002 than in 2001, April versus May. Thus, day length at the time of sea ice advance was about twice as long in 2002, 9 h versus 5 h, possibly one of the factors leading to the order of magnitude difference in the biomass of ice algae in the ice cores between the years (Table 2). Thus, in mid-winter when larval krill need to feed, ear- lier forming ice will have higher concentrations of ice algae than later forming ice. Ultimately with later forming ice, lower food concentrations of ice algae leads to less food available for the larval krill, lower growth rates, and lower predicted survivorship rates (Figure 3). We suggest that this mechanism underlies the correlation seen between the tim- ing of sea ice advance and recruitment success in the Palmer FIGURE 2. Growth rate (mm d H110021 ) of larval krill as a function of feeding index (pigment content, H9262g chl a g wwt H110021 ) of krill from same school. Growth rate is average for an experiment, with n H11005 3 to 14 individual measurements per experiment. Pigment content is average of 3 to 6 subsamples of larvae from same school, with 15? 20 larvae per subsample. Filled circles H11005 2001, open circles H11005 2002. Exponen- tial equation, r 2 H11005 0.81. FIGURE 3. Concept of mechanism underlying the correlation found between timing of ice advance and recruitment in Antarctic krill ( Siegel and Loeb, 1995; Quetin and Ross, 2003). 21_Quetin_pg285-298_Poles.indd 29321_Quetin_pg285-298_Poles.indd 293 11/17/08 8:47:57 AM11/17/08 8:47:57 AM 294 SMITHSONIAN AT THE POLES / QUETIN AND ROSS LTER study region (Quetin and Ross, 2003; Quetin et al., 2007). When ice does not form until May or June there is little in the water column to scavenge and PAR is near or at the minimum for the year. Thus, net primary production from ice formation to mid-winter will be low, and as a con- sequence so will food for larval krill. POTENTIAL IMPACT OF CLIMATE CHANGE The development of this conceptual view of the mechanism(s) underlying the correlation found between timing of sea ice advance and recruitment success in the Palmer LTER study region west of the Antarctic Peninsula underscores the importance of sea ice in the life cycle of Ant- arctic krill, and enhances our ability to predict how climate changes might impact krill population dynamics (Quetin et al., 2007). In a recent paper, Quetin et al. (2007) discuss the various scenarios and combinations of sea ice, light regime, and presence of Antarctic Circumpolar Deep Water that would create an optimal habitat for Antarctic krill. The Palmer LTER study region is situated in one of the fastest warming regions of the world, with the other two in the northern hemisphere, the Svalbard Island group and the Bering Sea. The evidence of warming west of the Antarctic Peninsula comes from multiple studies: the air temperatures are rapidly warming, with an increase in winter temperatures over the last 50 years of about 6?C (Smith and Stammerjohn, 2001; Vaughan et al., 2003); there is a warming of ocean temperatures at both surface and sub-surface depths (Gille, 2002; Meredith and King, 2005); and ice shelves and marine glaciers are retreating (Scambos et al., 2003; Cook et al., 2005). With the warm- ing climate, the duration of winter sea ice is shorter, but perhaps more importantly the timing is changing? sea ice advance is now later and retreat earlier (Parkinson, 2002; Smith et al., 2003; Stammerjohn et al., 2008). Sea ice ad- vance west of the Antarctic Peninsula is now usually in April or May, whereas in the late 1970s sea ice advanced in March (Parkinson, 2002). In a recent analysis of the 25- year satellite record for sea ice, Stammerjohn found that the mean day of advance is 20? 30 days later in the latter half of that period (1992? 2004) than in the earlier half (1979? 1991) (Stammerjohn et al., 2008). From the model simulations of Fritsen (1998), the impact of the 20- to 30- day delay in advance on accumulated SIMCO biomass in the sea ice between ice formation and mid-winter is likely to be substantial. Do we have any evidence of changes in the Antarctic krill population concurrent with this regional warming? Two studies to the north of the Palmer LTER study region suggest that populations of Antarctic krill are in decline. Atkinson et al. (2004) collated and compared trawl data from diverse studies in the Southern Ocean between 1926? 2003, and concluded that stocks of Antarctic krill in the southwest Atlantic have declined since the 1970s by a fac- tor of two. Shorter-term and smaller-space scale studies at the northern tip of the Antarctic Peninsula have included both net and bioacoustic data. The net data suggest a de- cline in krill stocks (Siegel, 2000) whereas the acoustic data suggest a cycle (Hewitt et al., 2003). One of the diffi culties in the analysis of these time-series data is that detecting a linear trend in data that exhibit a repetitive cycle will take many years. In the Palmer LTER study region, where we have shown that the pattern of episodic recruitment leads to a fi ve- to six-year cycle in the abundance of Antarctic krill (Quetin and Ross, 2003), a linear trend was not detectable in the 12-year time series (Ross et al., 2008). In this last sec- tion, we show the same data (methods and results in Quetin and Ross (2003) and Ross et al. (2008) from a different perspective, incorporating our understanding of the pre- dictability of the cycle in the population dynamics and in- terannual variability in the pattern of abundance. With the fi ve- to six-year cycle and two sequential years of successful recruitment followed by several years of low to no recruit- ment (Quetin and Ross 2003) in the LTER study region, the peak biomass in the cycle will appear in the January follow- ing the second good year class? in the time-series to date, in January of 1997 and 2003 (Figure 4). We can also look at the trend in the abundance during the fourth January: 1993, 1998 and 2004 (Figure 4). In both instances (year of maximum abundance, year 4 in cycle) the abundance FIGURE 4. Mean abundance of Antarctic krill within the Palmer LTER study region from 1993 to 2004, calculated with equations of the delta distribution as detailed in Ross et al. (2008). The dotted line follows the year of maximum abundance within the 5? 6 yr cycle, and the solid line follows the fourth year within the cycle. 21_Quetin_pg285-298_Poles.indd 29421_Quetin_pg285-298_Poles.indd 294 11/17/08 8:47:59 AM11/17/08 8:47:59 AM LIFE UNDER ANTARCTIC PACK ICE 295 has declined by 40%? 45% (Figure 4). We suggest that this analysis provides preliminary evidence from the Palmer LTER study region that populations of Antarctic krill are declining in this region in concert with the change in timing of advance of sea ice. However, to date, the analysis only encompasses two full cycles; an additional cycle may yield a different trend. SUMMARY Scuba diving research during the past 30 years has enhanced our understanding of the linkages between Ant- arctic krill and sea ice. We have been able to make key observations and conduct experiments on the dependency of larval Antarctic krill on the SIMCOs in the pack ice habitat. Our conceptual understanding of the ecology of the pack ice habitat and the intricacies of the interactions has greatly increased due to these activities. ACKNOWLEDGMENTS We would like to gratefully acknowledge the captains, crews, and support teams (from the NSF contractor, tech- nicians, and volunteers) that have made our research over the years both possible and enjoyable. Discussions with some of our colleagues have helped develop the concepts and ideas contained within this manuscript, and those from whom we have most benefi ted in recent years are C. H. Fritsen (Desert Research Institute), M. Vernet (Scripps Institution of Oceanography), and S. Stammerjohn (NASA Goddard Institute for Space Studies). This mate- rial is based upon work supported by the National Science Foundation, Offi ce of Polar Programs, under Award Nos. OPP-9011927 and OPP-9632763, and OPP-0217282 for the Palmer LTER, OPP-9909933 for Southern Ocean GLOBEC, and ANT-0529087 for PIIAK, The Regents of the University of California, the University of California at Santa Barbara, and the Marine Science Institute, UCSB. This is Palmer LTER contribution no. 315. NOTE 1. Since 1993 the Palmer LTER, a multidisciplinary program fo- cused on the pelagic ecosystem west of the Antarctic Peninsula (Smith 1995), has conducted research cruises in January/February, sampling a geographical area from the southern end of Anvers Island to Marguerite Bay to the south. The sampling grid is composed of fi ve transect lines ex- tending approximately 200 km offshore, with stations every 20 km, and 100 km apart alongshore, and covers an area of nearly 80,000 km 2 . LITERATURE CITED Atkinson, A., R. S. Shreeve, and A. G. Hirst. 2006. Natural Growth Rates in Antarctic Krill (Euphausia superba): II. 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CCAMLR Science, 6: 71? 84. 21_Quetin_pg285-298_Poles.indd 29821_Quetin_pg285-298_Poles.indd 298 11/17/08 8:48:01 AM11/17/08 8:48:01 AM ABSTRACT. The Ross Sea polynya is one of the most productive areas of the Southern Ocean; however, little is known about how plankton there respond to inhibitory solar exposure, particularly during the early-spring period of enhanced UVB (290? 320 nm) due to ozone depletion. Responses to solar exposure of the phytoplankton and bacterial assemblages were studied aboard the research ice breaker Nathaniel B. Palmer during cruises NBP0409 and NBP0508. Photosynthesis and bacterial production (thymidine and leucine incorporation) were measured during in situ incubations in the upper 10 m at three stations, which were occupied before, during, and after the annual peak of a phytoplankton bloom dominated by Phaeocystis antarctica. Near-surface production was consistently inhibited down to 5? 7 m, even when some surface ice was present. Relative inhibition of phytoplankton increased and productivity decreased with increas- ing severity of nutrient limitation as diagnosed using F v /F m , a measure of the maximum photosynthetic quantum yield. Relative inhibition of bacterial production was high for both the high-biomass and postbloom stations, but sensitivity of thymidine and leucine uptake differed between stations. These results provide the fi rst direct evidence that solar exposure, in particular solar ultraviolet radiation, causes signifi cant inhibition of Ross Sea productivity. INTRODUCTION Solar radiation, particularly that in the ultraviolet waveband (UV, 290? 400 nm), affects planktonic processes in the surface layer of diverse aquatic environ- ments (polar and elsewhere) and, in particular, the metabolism and survival of bacterioplankton, phytoplankton, and zooplankton. A subject of much recent work has been the extent to which these effects are augmented by enhanced UVB (290? 320 nm) due to Antarctic ozone depletion, which is most severe during the springtime ?ozone hole.? UVB-induced DNA damage has been measured in a wide variety of environments and trophic levels, for example, planktonic com- munities from tropical (Visser et al., 1999) and subtropical waters (Jeffrey et al., 1996a, 1996b), coral reefs (Lyons et al., 1998), and the Southern Ocean (Kel- ley et al., 1999; Buma et al., 2001; Meador et al., 2002). DNA damage in zoo- Patrick J. Neale, Jesse Phillips-Kress, and Linda A. Franklin, Smithsonian Environmental Research Center, 647 Contees Wharf Road, Edgewater, MD 21037, USA. Wade H. Jeffrey and J. Dean Pakulski, Center for Environmental Diagnostics and Bioremediation, University of West Florida, 11000 University Parkway, Building 58, Pen- sacola, FL 32514, USA. Cristina Sobrino, Smith- sonian Environmental Research Center; now at Departamento de Ecolog?a y Biolog?a Animal, University of Vigo, 36310 Vigo, Spain. Amy J. Baldwin, Center for Environmental Diagnostics and Bioremediation; now at Florida Department of Environmental Protection, 160 Governmental Center, Pensacola, FL 32502-5794, USA. Hae- Cheol Kim, Smithsonian Environmental Research Center; now at Harte Research Institute for Gulf of Mexico Studies, 6300 Ocean Drive, Corpus Christi, TX 78412, USA. Corresponding author: P. Neale (nealep@si.edu). Submitted 26 October 2007; revised 21 June 2008; accepted 28 May 2008. Inhibition of Phytoplankton and Bacterial Productivity by Solar Radiation in the Ross Sea Polynya Patrick J. Neale, Wade H. Jeffrey, Cristina Sobrino, J. Dean Pakulski, Jesse Phillips- Kress, Amy J. Baldwin, Linda A. Franklin, and Hae-Cheol Kim 22_Neale_pg299-308_Poles.indd 29922_Neale_pg299-308_Poles.indd 299 11/17/08 9:35:39 AM11/17/08 9:35:39 AM 300 SMITHSONIAN AT THE POLES / NEALE ET AL. plankton and fi sh larvae has been reported in the Southern Ocean (Malloy et al., 1997) and in anchovy eggs and larvae ( Vetter et al., 1999). The UV responses of Antarctic phytoplankton have been the focus of many studies (e.g., El-Sayed et al., 1990; Holm-Hansen and Mitchell, 1990; Mitchell, 1990; Helbling et al., 1992; Lubin et al., 1992; Smith et al., 1992; Boucher and Pr?zelin, 1996). However, there is little quantitative information on the photosynthetic response to UV in the Ross Sea and on the responses of natural assemblages of the colonial prymnesiophyte Phaeocystis antarctica, despite the important contribution of the Ross Sea to overall pro- ductivity of the Southern Ocean (see Smith and Comiso, 2009, and references therein). P. antarctica is the domi- nant phytoplankter in the Ross Sea, particularly during the early-spring period of ozone depletion. At this time of year most of the Ross Sea is covered by ice, so phytoplankton growth occurs in an open water area, or polynya, located just north of the Ross Ice Shelf (for more background, see DiTullio and Dunbar, 2004). Our lack of knowledge about responses to UV is not only for P. antarctica but also for other phytoplankton and the associated bacterioplankton community. Bacterioplankton abundance can reach 3 H11003 10 9 cells/L in the Ross Sea, equal to bacterial blooms in other oceanic systems. Bacterioplankton do bloom in response to the Phaeocystis bloom, but with a delay of one or two months after the onset of the phytoplankton bloom (Ducklow et al., 2001). DOC release by Phaeocystis is low, but is be- lieved to be labile (Carlson et al., 1998) and may limit bac- terial production in the upper water layer (Ducklow et al., 2001). Bacterial production in deeper waters is relatively high (Ducklow et al., 2001) and may be related to sinking Phaeocystis POC (DiTullio et al., 2000). There are many other measurements to suggest that enhanced UVB and environmental UV in general have ef- fects on organismal physiology and survival (reviewed in de Mora et al., 2000). Direct measurements of quantitative in situ effects, on the other hand, are diffi cult to make for most cases. However, estimates can be made using mathematical models. The quantitative response to UV exposure is char- acterized well enough for some processes that statements can be made about integrated effects over the water column as a function of vertical mixing in the surface layer (Neale et al., 1998; Huot et al., 2000; Kuhn et al., 2000). These model results, together with profi les of UV-specifi c effects like DNA damage under qualitatively different mixing con- ditions (Jeffrey et al., 1996b; Huot et al., 2000), argue that mixing signifi cantly modifi es water column effects (Neale et al., 2003). However, there are no instances where UV re- sponses and vertical mixing have been quantitatively mea- sured at the same time. Here we present results from fi eld work conducted in the Ross Sea polynya to assess the quantitative impact of UV on the phytoplankton and bacterioplankton com- munities. Both communities play a crucial role in carbon and nutrient cycling. They are also tightly coupled, so it is important to examine both communities simultaneously to understand UV impacts on the system as a whole. For example, a decrease in phytoplankton production may result in a decline in bacterial production that may be compounded by direct UVB effects on bacterioplankton. A primary physical factor controlling exposure of these communities to UV is vertical mixing. Thus, our work ex- amined the effects of vertical mixing using a combination of fi eld measurements and modeling approaches. Our assessments of UV responses of Ross Sea plank- ton used three approaches: laboratory spectral incubations, surface (on deck) time series studies, and daylong in situ incubations. The fi rst two approaches enable estimation of spectral response (biological weighting functions, Cullen and Neale, 1997) and kinetic response. From this informa- tion we are constructing general, time-dependent models of UV response to variable irradiance in the mixed layer. While providing less detail on specifi c responses, in situ in- cubations have the advantage of using natural irradiance regimes. However, they are not suffi cient in themselves in measuring actual water column effects since they intro- duce the artifact of keeping samples at a constant depth throughout the day. For example, depending on the kinet- ics of UV inhibition, a static incubation may overestimate the response at the surface but underestimate the integrated response over the water column (Neale et al., 1998). Nevertheless, in situ incubations still provide useful in- formation on responses of natural plankton assemblages. They provide direct evidence that UV exposure is suffi - ciently high to cause some effect, in particular, inhibition of near-surface productivity. Moreover, in situ observations can be compared to predictions of laboratory-formulated models evaluated using measured irradiance at the incuba- tion depths and thus provide an independent validation of the models. Here we present measurements of phytoplankton productivity ( 14 C-HCO3 H11002 incorporation) and bacterial production ( 3 H-leucine and 3 H-thymidine incorporation) for daylong incubations conducted in the near surface (upper 10 m) of the Ross Sea polynya for three dates spanning the early-spring through summer period. 22_Neale_pg299-308_Poles.indd 30022_Neale_pg299-308_Poles.indd 300 11/17/08 9:35:39 AM11/17/08 9:35:39 AM INHIBITION OF PHYTOPLANKTON AND BACTERIAL PRODUCTIVITY 301 MATERIALS AND METHODS IRRADIANCE MEASUREMENTS Radiometers were mounted on top of a science mast (nominally 33 m above ocean surface). Photosynthetically available radiation (PAR, 400? 700 nm) incident on a fl at plane (2-H9266 collector) was measured with a Biospherical Instruments (San Diego, California, USA) GUV 2511. Spectral UV irradiance was recorded with a Smithsonian- designed multifi lter radiometer, the SR19, which measures between 290 and 324 nm with 2-nm bandwidth (FWHM) and resolution and at 330 nm with 10-nm bandwidth (technical description in Lantz et al., 2002). Broadband UV measurements (nominal 10 nm bandwidth) in the UV were also made by the GUV 2511. The transmission of downwelling irradiance (E d [?]) through the water column was measured by a free-fall, profi ling radiometer, the Bio- spherical Instruments PUV 2500. Four to fi ve casts were made near solar noon from 0 to 50 m, and attenuation coeffi cients (k d [?]) were computed from the regression of log(E d [?]) versus depth. Profi les of E d were recorded at 305, 313, 320, 340, and 395 nm (only k d [?] are presented here) and for PAR. PRODUCTIVITY ASSAYS Sample water for the incubations was obtained with 30-L ?Go-Flo? Niskin bottles (General Oceanics) mounted on a conductivity-temperature-depth (CTD) rosette. The CTD cast was made in open water at 5-m depth at approxi- mately 0500 local time (LT) (GMTH1100113), ensuring minimal exposure to UV prior to incubation. The sample was im- mediately dispensed through wide-bore tubing and stored in the dark at 0?C until use. For photosynthesis assays, UV- transparent polyethylene sample bags (113-mL Whirl-Pak bag) were prepared by extensive rinsing with sample wa- ter. Then 14 C-bicarbonate was added to 700 mL of sample water (H110111 H9262Ci/mL), which was distributed into 14 sample bags in 50-mL aliquots. The unfi lled portion of the bag was tightly rolled and twist sealed to prevent leakage. A second set of bags was prepared for measurements of bacterial pro- ductivity. Tritium ( 3 H) labeled thymidine (60 Ci/mmol) or 3 H-leucine (60 Ci/mmol) was added to 175 mL of seawater to a fi nal concentration of 10 nM for 19 January and 21 November and 20 nM for 28 November. Five milliliters of the amended solution were added to each of three Whirl- Pak bags, as for photosynthesis, such that triplicates for each substrate were placed at each depth. After inoculation, the bags for both photosynthesis and bacterial productiv- ity were secured with plastic ties to 25 H11003 25 cm ?crosses? made of UV-transparent acrylic sheet (Plexiglas) (Figure 1). Each ?arm? was 10 cm wide; one set of replicate photo- synthesis bags was fastened to one set of opposing arms, and triplicate 3 H-thymidine incorporation and triplicate 3 H-leucine incorporation bags were attached to each of the other two arms. Crosses were kept at 0?C and in the dark until just before deployment. These Plexiglas pieces were then secured at 1-, 2-, 3-, 4-, 5-, 7.5- and 10-m depth to a weighted line which passed through the center of each cross. A primary fl oat was attached at the surface along with a second fl oat containing a radar refl ector and a radio beacon. The array was hand deployed from the stern of the research vessel and followed for 12 h. Upon retrieval of the array, bags were quickly removed from the arms and transported to the laboratory in the dark. For photosyn- thesis, fi ve replicate aliquots (5 mL) were removed from the bags and analyzed for incorporated organic 14 C-carbon by acidifi cation, venting and scintillation counting. Replicate 1.5-mL samples for 3 H-thymidine or 3 H-leucine incorpora- tion were removed from each Whirl-Pak bag and placed in 2-mL microfuge tubes containing 100 H9262L of 100% tri- chloroacetic acid (TCA). Samples were processed via the FIGURE 1. Schematic diagram of ?cross? supports for the in situ array. The cross pieces are 25 cm in length. Darker shaded boxes indicate position of the UV-transparent polyethylene (Whirl-Pak) in- cubation bags. The center circle indicates where the support attached to the incubation line. 22_Neale_pg299-308_Poles.indd 30122_Neale_pg299-308_Poles.indd 301 11/17/08 9:35:40 AM11/17/08 9:35:40 AM 302 SMITHSONIAN AT THE POLES / NEALE ET AL. microcentrifugation method described by Smith and Azam (1992) as modifi ed by Pakulski et al. (2007). BIOMASS AND PHOTOSYNTHETIC QUANTUM YIELD Chlorophyll concentration and bacterial cell abundance was measured on aliquots of the early-morning 5-m sample at all cruise stations (including the incubation stations). Samples for chlorophyll were concentrated on glass fi ber fi l- ters (GF/F, Whatman Inc., Florham Park, New Jersey, USA) and extracted with 90% acetone overnight at 0?C. After extraction, chlorophyll concentration was measured as the fl uorescence emission in a Turner 10-AU fl uoro meter cali- brated with pure chlorophyll a (Sigma Chemical, St. Louis, Missouri, USA). Bacterial abundances were determined by epifl uorescence microscopy of 5? 10 mL of 4H11032, 6-diamidino- 2-phenylindole (DAPI) stained samples collected on black 0.2-H9262m polycarbonate fi lters (Porter and Feig, 1980). A pulse-amplitude-modulated fl uoro meter (Walz Water PAM, Effeltrich, Germany) with red LED (650 nm) excitation was used to assess the maximum photosynthetic effi ciency (quantum yield) of the samples. Measurements on the 5-m sample were made after at least one hour of dark incuba- tion at 0?C. The data are expressed as the PSII quantum yield, F v /F m H11005 (F m -F 0 )/F m , which has been correlated with the maximum quantum yield of photosynthesis (Genty et al., 1989). F 0 is the steady-state yield of in vivo chlorophyll fl uorescence in dark-adapted phytoplankton, and F m is the maximum yield of fl uorescence obtained from the same sample during application of a saturating light pulse (400- ms duration). SITE DESCRIPTION The Ross Sea polynya was sampled in two research cruises aboard the R/V Nathaniel B. Palmer taking place in December 2004 to January 2005 (NBP0409) and October through November 2005 (NBP0508). Overall trends in surface biomass are shown in Figure 2. During both years, this south central region of the Ross Sea supported a strong bloom of P. antarctica in November, peaking in early December (on the basis of Moderate Resolution Im- aging Spectroradiometer (MODIS) satellite images). The bloom slowly declined through January, becoming mixed with other species, mostly diatoms. Bacterial biomass dis- played a more complex pattern, with biomass peaks occur- ring during each of the cruise periods. Bacterioplankton FIGURE 2. Time series of chlorophyll and bacterial cell concentration at 5 m for all stations in two cruises to the Ross Sea polynya. Bars indi- cate standard deviation of triplicate determinations. The two sampling periods for NBP0409 (December 2004 to January 2005) and NBP0508 (November 2005) are indicated by horizontal lines, and vertical arrows indicate dates of incubations. 22_Neale_pg299-308_Poles.indd 30222_Neale_pg299-308_Poles.indd 302 11/17/08 9:35:52 AM11/17/08 9:35:52 AM INHIBITION OF PHYTOPLANKTON AND BACTERIAL PRODUCTIVITY 303 are seen to increase along with the onset of the bloom followed by a second peak occurring in mid-January as the bloom receded. Our data from October? November is very similar to Ducklow et al. (2001), but this previous study and ours differ for the December? January period. We observed relatively low bacterial numbers at the end of December when the cruise began. Bacterioplankton then increased to a second peak occurring at approximately the same time as that reported by Ducklow et al. (2001) but at a maximum density of only 0.6 H11003 10 9 cells/L compared to the H110112 H11003 10 9 cells/L reported in the previous study. These contrasting observations may be due to differences in spe- cifi c bloom conditions between years or specifi c sampling locations within the Ross Sea. Deployment locations and times of the incubations are given in Table 1. During the early-spring (October? November) cruise, the surface was covered with moderate to heavy pack ice interspersed with leads until the last week in November. For the fi rst incuba- tion (21 November), samples were obtained and the array was deployed while the ship was in a lead. Shortly after deployment, the array became surrounded with a raft of ?pancake? ice extending at least a 100 m in all directions (Figure 3), and this continued until retrieval. The 28 No- vember and 19 January deployments were conducted in open water. RESULTS SOLAR IRRADIANCE Surface UV and PAR were similar between all three days, with midday PAR in the range of 1000? 1200 H9262mol m H110022 s H110021 and midday UV at 320 nm between 100 and 150 mW m H110022 nm H110021 (Figure 4). Transmission of UV and PAR varied between dates in inverse relation to phytoplank- ton biomass. Attenuation coeffi cients were similar for the prebloom and postbloom stations but were considerably higher in both UV and PAR for the station on 28 Novem- ber near the peak of the bloom (Table 1). PHOTOSYNTHESIS All in situ profi les exhibited lowest rates at the surface and higher rates with depth, with the near-surface ?photo- active? zone of inhibitory effect extending to at least 5 m on all dates (Figure 5). The 21 November profi le shows an inhibitory trend over the full profi le, but differences below 4 m are not signifi cant due to high sample variability. This high variability may be associated with ice-cover-generated heterogeneity in the underwater light fi eld. Interestingly, relative inhibition at 1 m is only 10% less than in the 28 November profi le, despite the presence of ice cover on 21 November (Figure 2). On 28 November, the depth maxi- mum in productivity was observed at 5 m, which was much shallower than the other dates. This is consistent with the relatively low transparency to both PAR and UV on this date due to high phytoplankton biomass (5.5 mg m H110023 ), mostly FIGURE 3. Typical surface conditions during the 21 November in- cubation. The surface fl oat of the array sitting on top of the ice is approximately 75 cm in diameter. TABLE 1. Background information on the three stations where in situ incubations were conducted. LT H11005 local time. Date (LT) Latitude Longitude Chl a (mg m H110023 ) k d [320] (m H110021 ) k d PAR (m H110021 ) 21 Nov 2005 H1100277?35.113H11032 178?23.435H11032 1.9 0.32 0.15 28 Nov 2005 H1100277?34.213H11032 H11002178?57.763H11032 5.5 0.54 0.27 19 Jan 2005 H1100274?30.033H11032 173?30.085H11032 2.8 0.32 0.17 22_Neale_pg299-308_Poles.indd 30322_Neale_pg299-308_Poles.indd 303 11/17/08 9:35:55 AM11/17/08 9:35:55 AM 304 SMITHSONIAN AT THE POLES / NEALE ET AL. comprised of P. antarctica (data not shown). The presence of P. antarctica decreased not only PAR transparency be- cause of absorbance by photosynthetic pigments but also UV transparency (Table 1). The decreased UV transparency is caused in part by the presence of UV screening pigments, the mycosporine-like amino acids, which are known to be accumulated by this species and were present in separate absorbance scans of particulates (data not shown). The 19 January incubation was at a postbloom station for which the depth of UV effects is comparable to the prebloom 21 November station and relative inhibition at 1 m was the highest of all profi les. If the three profi les are regarded as showing the se- quential development of the Ross Sea polynya bloom (despite the 19 January station being from the previous season), a couple of trends are apparent. One is the large increase in productivity associated with the high bio- mass on 28 November. Also, productivity was higher in the prebloom station compared to the postbloom station despite similar biomass. In other words, biomass-specifi c maximum productivity in the profi le (P B max , at 5 m on 28 November and 10 m on 21 November and 19 January) was highest before the bloom and actually decreased with time (Figure 6). Parallel to this result was a decrease in the maximum quantum yield of photosynthesis as measured by PAM fl uorometry (Figure 6). The most likely reason for the declining quantum yield, which has been observed pre- viously for postbloom phytoplankton in the Ross Sea (e.g., Peloquin and Smith, 2007), is the depletion of dissolved iron, the limiting nutrient for phytoplankton growth in most areas of the Ross Sea (Smith et al., 2000). An ad- ditional factor could be the cumulative effect of recurring inhibitory solar exposure on the functioning of the photo- synthetic apparatus. BACTERIOPLANKTON PRODUCTION Similar to the pattern observed for photosynthetic rates, bacterial incorporation of either leucine or thymidine was most inhibited at the high-biomass and postbloom stations. For leucine incorporation, the lowest rates and least inhibition at 1 m were observed for the early-season sample, while the highest production rates were observed in the high-biomass sample. The pattern was similar for thymidine incorporation, although there was minimal dif- ference between the high-biomass and postbloom samples. The pattern of dark leucine rates generally followed bacte- rial biomass (Figure 2), with minor variation in rates per cell (not shown). In contrast, thymidine rates remained high at the postbloom station. DISCUSSION The results presented here show some of the fi rst observations of the effects of full-spectrum, near-surface solar exposure on plankton assemblages in the Ross Sea polynya from which we can already make several conclu- FIGURE 4. Daily variation in the surface quantum fl ux of photosynthetically available radiation (H9262mol m H110022 s H110021 PAR, 400? 700 nm, solid line) and spectral irradiance at 320 nm (mW m H110022 nm H110021 , 2.0-nm bandwidth at half maximum, dashed line). Vertical lines indicate period of incuba- tion on each date. 22_Neale_pg299-308_Poles.indd 30422_Neale_pg299-308_Poles.indd 304 11/17/08 9:35:59 AM11/17/08 9:35:59 AM INHIBITION OF PHYTOPLANKTON AND BACTERIAL PRODUCTIVITY 305 sions. First, it is clear that incident solar exposure is suf- fi ciently high and plankton assemblages are suffi ciently sensitive that inhibition of near-surface algal and bacterial productivity is a regular occurrence during the spring? summer period in the southern Ross Sea. Photosynthesis was more strongly inhibited than bacterial productivity, such that effects on phytoplankton could even be observed below light ice cover (21 November). These effects were also observed even though UVB exposure was not signifi - cantly enhanced by ozone depletion. Although low ozone can occur throughout November in the Ross Sea region (Bernhard et al., 2006), the ?ozone hole? was not present during the NBP0508 incubations. Indeed, UV and PAR exposure in the Ross Sea polynya were not high compared to other observations in Antarctic waters (Kieber et al., 2007; Pakulski et al., 2007). Thus, the assemblages must be particularly sensitive to solar ex- posure in order for effects to be so pronounced despite moderate exposure levels. This conclusion is consistent with the preliminary results of our laboratory measure- ments of biological weighting functions for UV inhibition. These showed the highest sensitivity to UV yet recorded for Antarctic phytoplankton and modest sensitivity to UV for Ross Sea bacterioplankton (Neale et al., 2005; Jeffrey et al., 2006). They also showed that most of the inhibitory effect of near-surface irradiance on photosynthesis was FIGURE 5. Hourly productivity rates for photosynthesis (P) and bacterial incorporation of leucine (Leu) and thymidine (Tdr) for the incuba- tions on 21 November, 28 November, and 19 January (all in 2005). The bottom symbols for Leu and Tdr show rates for samples incubated in the dark. Horizontal bars indicate assay standard deviation (P, n H11005 10; Leu/Tdr, n H11005 6). The numbers below each profi le show the percent inhibition at 1 m relative to the peak rate in the profi le (photosynthesis) or rate in the dark (bacterial incorporation). FIGURE 6. Measurements on the early-morning 5-m sample used for the in situ incubations. Maximum chlorophyll-specifi c rate of photosynthesis of the in situ incubation (black bars, left axis) and maximum photosynthetic quantum yield as measured with PAM fl uorometry (gray bars, right axis). These are two independent ap- proaches to indicate the relative variation in the overall photosyn- thetic capacity of the sampled phytoplankton assemblage. 22_Neale_pg299-308_Poles.indd 30522_Neale_pg299-308_Poles.indd 305 11/17/08 9:36:57 AM11/17/08 9:36:57 AM 306 SMITHSONIAN AT THE POLES / NEALE ET AL. due to UV, with PAR having only a small effect. Similarly, Smith et al. (2000) did not observe signifi cant near- surface inhibition when they measured daily in situ primary pro- ductivity in the Ross Sea using UV-opaque enclosures, although PAR inhibition was observed in on-deck incuba- tions receiving higher than in situ irradiance. This high sensitivity to UV may be a consequence of the acclimation to low-irradiance conditions in the early-season assem- blage (before iron depletion) and iron limitation during the late season. The lower sensitivity of bacterioplankton to UVR may have been related to nutrient replete condi- tions. Three separate experiments over the course of the sampling period during November 2005 indicated that the bacterioplankton were not nutrient (Fe, N, C) lim- ited (data not shown). Our previous work has suggested that alleviation of nutrient limitation often reduced UVR sensitivity (Jeffrey et al., 2003). Unfortunately, no data is available for the summer 2005 samples. Bacterioplankton abundance increased as did chl a during this period, in contrast to the lags reported by others. Our observation may be, in part, due to the apparent replete nutrient con- ditions we observed. Although Ducklow et al. (2001) re- ported low DOC production by the Phaeocystis bloom, it is labile (Carlson et al., 1998) and it has been hypothesized that macronutrient depletion seldom occurs in the Ross Sea (Ducklow et al., 2001). Results have been combined for two years; however, the time course of phytoplankton biomass in the Ross Sea for both 2004? 2005 and 2005? 2006 followed the normal pattern of peak biomass at the end of November (Peloquin and Smith, 2007). Although species composition shifted be- tween the cruises, inhibition was consistently high for all profi les. In contrast, bacterial response was less consistent between the cruises. The ratio between leucine and thymi- dine dark uptake was H1102210 in November 2005 but H1102110 in January 2005, suggesting basic metabolic differences between assemblages. The abundance patterns during the cruises also show separate growth ?events? occurring dur- ing each cruise (though some of the variation may be due to spatial differences). These observations suggest that the two cruises sampled physiologically distinct bacterial as- semblages, a conclusion that is consistent with differences in sample genetic composition as determined using terminal restriction fragment length polymorphisms (TRFLP) analy- sis (A. Baldwin, University of West Florida, and W. H. Jef- frey, University of West Florida, personal communication). In summary, our results provide direct evidence that in situ UV irradiance in the Ross Sea is inhibitory for both phytoplankton photosynthesis and bacterioplankton production. In terms of the magnitude of the responses observed in the incubations, these should be conservative estimates of the effects of solar exposure on in situ plank- tonic production. Models of UV- and PAR-dependent photosynthetic response, when evaluated for the exposure occurring at each depth in the array, predict a compara- ble response as observed in situ (Neale et al., 2005). In contrast, vertical profi les of fl uorescence-based photosyn- thetic quantum yield showed that inhibited phytoplank- ton are found deeper in the water column than the 5-7 m depth of the photoactive zone in the incubations. This en- hancement of inhibition in the water column is consistent with vertical exchange due to both Langmuir ciculation and near-surface internal waves, both of which increase the proportion of surface layer phytoplankton exposed to inhibiting irradiance. The operation of these mechanisms was confi rmed by physical measurements. Detailed com- parisons of production estimates using these multiple ap- proaches will be presented in subsequent reports. ACKNOWLEDGMENTS We thank the captain and crew of the R/V Nathaniel B. Palmer and Raytheon Polar Services Co. for the fi eld support provided. The authors gratefully acknowledge Ronald Kiene, Dauphin Island Sea Lab, Chief Scientist on NBP0409, for providing the PAM fl uorometer, and Hyakubun Harada, Dauphin Island Sea Lab, for assis- tance in its use. Support was provided by National Science Foundation Offi ce of Polar Programs grant 0127037 to PJN and 0127022 to WHJ. CS was supported by an As- turias Fellowship from the Scientifi c Committee for Ant- arctic Research (SCAR) and by the Spanish Ministry of Education and Science (MEC). LITERATURE CITED Bernhard, G., C. R. Booth, J. C. Ehramjian, and S. E. Nichol. 2006. 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In Phaeocystis, Major Link in the Biogeochemical Cycling of Elements, p. 16. SCOR Working Group, No. 120. Groeningen, Netherlands: SCOR Working Group. Pakulski, J. D., J. A. Meador, J. P. Kase, and W. H. Jeffrey. 2007. Effect of Stratospheric Ozone Depletion and Enhanced Ultraviolet Radia- tion on Marine Bacteria at Palmer Station, Antarctica in the Early Austral Spring. Photochemistry and Photobiology, 83: 1? 7. Peloquin, J. A., and W. O. Smith. 2007. Phytoplankton Blooms in the Ross Sea, Antarctica: Interannual Variability in Magnitude, Tem- poral Patterns, and Composition. Journal of Geophysical Research, 112: C08013, doi: 10.1029/2006JC003816. Porter, K. G., and Y. S. Feig. 1980. The Use of DAPI for Identifying and Counting Aquatic Microfl ora. Limnology and Oceanography, 25: 243? 248. Smith, D. C., and F. A. Azam. 1992. A Simple, Economical Method for Measuring Bacterial Protein Synthesis Rates in Seawater Using 3h- Leucine. Marine Microbial Food Webs, 6: 107? 114. Smith, R. C., B. B. Pr?zelin, K. S. Baker, R. R. Bidigare, N. P. Boucher, T. Coley, D. Karentz, S. MacIntyre, H. A. Matlick, D. Menzies, M. Ondrusek, Z. Wan, and K. J. Waters. 1992. Ozone Depletion: 22_Neale_pg299-308_Poles.indd 30722_Neale_pg299-308_Poles.indd 307 11/17/08 9:37:59 AM11/17/08 9:37:59 AM 308 SMITHSONIAN AT THE POLES / NEALE ET AL. Ultraviolet Radiation and Phytoplankton Biology in Antarctic Wa- ters. Science, 255: 952? 959. Smith, W. O., Jr., and J. C. Comiso. 2009. ?Southern Ocean Primary Productivity: Variability and a View to the Future.? In Smithsonian at the Poles: Contributions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 309? 318. Washington, D.C.: Smithsonian Institution Scholarly Press. Smith, W. O., Jr., J. Marra, M. R. Hiscock, and R. T. Barber. 2000. The Seasonal Cycle of Phytoplankton Biomass and Primary Productiv- ity in the Ross Sea, Antarctica. Deep-Sea Research, Part II, 47: 3119? 3140. Vetter, R. D., A. Kurtzman, and T. Mori. 1999. Diel Cycles of DNA Dam- age and Repair in Eggs and Larvae of Northern Anchovy, Engraulis mordax, Exposed to Solar Ultraviolet Radiation. Photochemistry and Photobiology, 69: 27? 33. Visser, P. M., E. Snelder, A. J. Kop, P. Boelen, A. G. J. Buma, and F. C. van Duyl. 1999. Effects of UV Radiation on DNA Photodamage and Production in Bacterioplankton in the Coastal Caribbean Sea. Aquatic Microbial Ecology, 20: 49? 58. 22_Neale_pg299-308_Poles.indd 30822_Neale_pg299-308_Poles.indd 308 11/17/08 9:38:00 AM11/17/08 9:38:00 AM ABSTRACT. The primary productivity of the Southern Ocean south of 58?S is assessed using satellite data on ice concentrations, sea surface temperatures, and pigment con- centrations, a vertically generalized production model, and modeled photosynthetically active radiation. Daily productivity is integrated by month and by year to provide an estimate of new production. The productivity of the Southern Ocean is extremely low relative to other oceanic regions, with annual net rates throughout the region of less than 10 g C m H110022 . This low annual value is largely the result of negligible productivity throughout much of the year due to low irradiance and high ice cover. Despite the annual oligotrophic state, monthly productivity during the summer (December through Febru- ary) is substantially greater, averaging from 100 to 1,500 mg C m H110022 mo H110021 . Substantial interannual variability occurs, and certain subregions within the Southern Ocean experi- ence greater interannual variations than others. Those regions, like the West Antarctic Peninsula, the Ross Sea polynya region, and the Weddell Sea, are characterized as being continental shelf regions and/or those that are substantially impacted by ice. Despite this relationship, no signifi cant changes in primary production were observed in regions where large trends in ice concentrations have been noted. The driving forces for this vari- ability as well as the implications for long-term changes in regional and Southern Ocean productivity are discussed. INTRODUCTION The Southern Ocean is a vast region within the world?s oceans that has pre- sented some signifi cant challenges to oceanographers. It is the site of large num- bers of birds, marine mammals, and fi shes and extensive sedimentary deposits of biogenic material, and is presently being impacted by physical forcing external to the region, such as ozone depletion (Neale et al., 1998, 2009, this volume) and climate change (e.g., Vaughan et al., 2003). However, because of its size and remoteness, it is diffi cult to conduct experimental programs to adequately assess the role of various environmental factors on biological processes in the region. In addition, a large fraction of the Southern Ocean is ice covered for much of the year, restricting access to many locations and making sampling of other regions nearly impossible. To assess the productivity of the entire Southern Ocean, it is necessary to ?sample? using techniques that can quantify processes over large Walker O. Smith Jr., Virginia Institute of Marine Sciences, College of William and Mary, Glouces- ter Point, VA 23062, USA. Josefi no C. Comiso, NASA Goddard Space Flight Center, Code 614.1, Greenbelt, MD 27701, USA. Corresponding au- thor: W. O. Smith (wos@vims.edu). Accepted 28 May 2008. Southern Ocean Primary Productivity: Variability and a View to the Future Walker O. Smith Jr. and Josefi no C. Comiso 23_Smith_pg309-318_Poles.indd 30923_Smith_pg309-318_Poles.indd 309 11/17/08 8:44:17 AM11/17/08 8:44:17 AM 310 SMITHSONIAN AT THE POLES / SMITH AND COMISO spatial scales through time. At present, the only means to accomplish this on the appropriate scales is via satellite oceanography. Satellites presently have the capability to accurately map the distributions of ice (Comiso, 2004), sea surface temperature (SST; Comiso, 2000; Kwok and Comiso, 2002), and pigment concentrations (Moore and Abbott, 2000), as well as other parameters such as winds, ba- thymetry, cloud cover, and some gas concentrations such as ozone (Comiso, 2009). Some measurements use vis- ible wavelengths and refl ectance from the surface, and therefore the data returned are reduced in space and time because of clouds; others are either passively detected or use other wavelengths to determine the distribution of the variable. In biological oceanography a major variable of interest is ocean color, which is converted into quantitative estimates of pigment (chlorophyll) concentrations. While the estimates include signifi cant error terms (because of the dependence of pigment estimates as a function of lati- tude, the limitation of refl ectance to the optical surface layer rather than the entire euphotic zone, and the inter- ference in some waters of dissolved organic matter), these estimates remain, and will remain, the only means to ob- tain synoptic assessments of phytoplankton distributions over large areas as well as their temporal changes over relatively short (e.g., days) periods. Two satellites have provided nearly all of the data in the past three decades on pigment distributions in the South- ern Ocean. The fi rst was the Nimbus 7 satellite, launched in 1978, which carried the Coastal Zone Color Scanner (CZCS). While questions concerning the data quality and coverage from CZCS have been voiced, the data were used to investigate both the large-scale distributions of pigments in relation to oceanographic variables (Sullivan et al., 1993; Comiso et al., 1993) and also the specifi c processes and regions (e.g., Arrigo and McClain, 1994). However, given the orbit, the frequency of data collection in the Southern Ocean was quite restricted, and when compounded by the loss of data from cloud cover, the temporal frequency was far from optimal. In 1996 the ORBView-2 satellite was launched, which included the Sea-viewing Wide Field-of- view Sensor (SeaWiFS). This satellite proved to be an ex- tremely useful tool for biological oceanographers, as the sampling frequency was much greater and the data return in polar regions was far greater. For example, Moore et al. (1999) were able to detect a short-lived bloom in the Pacifi c sector of the Southern Ocean that was only infrequently sampled by ships. Dierssen et al. (2002) assessed the vari- ability of productivity in the West Antarctic Peninsula region and found (based on a model) that pigment concentrations were the dominant variable creating variations in space and time. Smith and Comiso (2008) assessed the productivity of the entire Southern Ocean and found that the ?hot spots? of production were limited to continental shelf regions, and suggested that this was a result of low iron concentra- tions coupled with deeper mixing in the offshore regions. The interaction of low iron and low irradiance (Sunda and Huntsman, 1997; Boyd and Abraham, 2001) gives rise to a large spatial limitation over broad areas. It is the purpose of this manuscript to look at the scales of variability in the Southern Ocean as a whole and to determine where such variations are large by using pri- mary production derived from SeaWiFS ocean color and advanced very high resolution radiometer (AVHRR) SST data in conjunction with a bio-optical model. We also will compare the modeled productivity with observed values, where those data are available to test the robustness of the model. Finally, some aspects of the temporal patterns of productivity in the Southern Ocean are reviewed. MATERIALS AND METHODS Primary productivity was estimated using various data derived from satellites and a bio-optical model. The model was a vertically generalized production model (Behrenfeld and Falkowski, 1997b), in which primary productivity (PP eu , in units of mg C m H110022 d H110021 ) was estimated from the following equation: PP P E E CZD eu opt B o o Sat eu Irr =? + ??0 66125 41 . . where P opt B is the optimal rate of photosynthesis within the water column (mg C (mg chl) H110021 h H110021 ) and is a function of temperature, E o is the surface daily photosynthetically active radiation (PAR, mol photons m H110022 d H110021 ), C sat is the surface chlorophyll concentration (mg chl m H110023 ) deter- mined by satellite, Z eu is the depth of the euphotic zone (m), and D Irr is the photoperiod (h). P opt B was estimated from sea surface temperatures by the polynomial equation of Behrenfeld and Falkowski (1997b), and all P opt B values at temperatures less than H110021.0?C were set to 1.13. Temperature, PAR, ice concentrations, and chloro- phyll concentrations were derived from different satel- lite data sets. Different satellite data were mapped to the same grid as described below. We arbitrarily defi ned the Southern Ocean roughly as the region impacted by sea- sonal ice movements and hence set the northern bound- 23_Smith_pg309-318_Poles.indd 31023_Smith_pg309-318_Poles.indd 310 11/17/08 8:44:17 AM11/17/08 8:44:17 AM SOUTHERN OCEAN PRIMARY PRODUCTIVITY 311 ary at 58?S. Ice concentrations and associated parameters (e.g., ice extent and area) were derived using data from the Special Sensor Microwave Imager (SSM/I) on the Defense Meteorological Satellite Program and mapped on a polar stereographic grid at a 25 H11003 25 km resolution. Ice con- centrations were derived from satellite passive microwave data using the enhanced bootstrap algorithm used for Ad- vanced Microwave Scanning Radiometer-EOS data and adapted for SSM/I data (e.g., Comiso et al., 2003; Comiso, 2004). Sea surface temperatures were derived from ther- mal infrared channels of the NOAA AVHRR as described in Comiso (2003). Pigment concentrations derived from SeaWiFS data were provided by the NASA Goddard Earth Sciences Distributed Active Archive Center. Surface tem- perature and pigment concentration data have been grid- ded in the same manner as the sea ice concentration data but on a 6.25 H11003 6.25 km resolution. Mean daily pigment concentrations were estimated using the standard SeaWiFS algorithm with OC4 (Version 4) calibration (Patt et al., 2003) and used to generate weekly (seven-day bins) and monthly data sets from 1997 to 2006. Photosynthetically active radiation data were extracted as part of the Sea- WiFS data but were not used in the estimates of produc- tivity because a large fraction of the valuable polar data was inadvertently masked as ice covered by the SeaWiFS data processing group. We used a modeled PAR instead (which provided basically the same results) for much im- proved coverage. It is important to recognize that because of cloud and ice masking the weekly and monthly averages do not refl ect true averages but are averages of daylight data (for each data element) available during clear-sky, ice-free conditions only. Productivity was calculated on a daily basis and binned in a manner similar to that of chlorophyll. The gridding technique (Smith and Comiso, 2008) and the presence of clouds caused a large fraction of data elements (pixels) in the daily maps to have missing data. In the case where an empty pixel is surrounded by pixels with data, a simple interpolation technique is utilized to estimate the pigment level in the empty pixel. For larger data gaps, a combination of spatial and temporal interpolation was utilized. Such interpolation fi lled only a very small fraction of missing data in the daily maps, and for time series stud- ies, weekly averages were produced as the basic product. In a similar manner, annual productivity was estimated by summing weekly averages over an entire year. Standard deviations were calculated for all pixels, but because of the variable number of data points within each pixel, we arbi- trarily used only those locations where at least fi ve means were available to calculate variations. We recognize that regional algorithms have been de- veloped for certain parts of the Southern Ocean (e.g., Ross Sea: Arrigo et al., 1998; Dierssen and Smith, 2000) and that these formulations provide a more accurate estimate of phytoplankton biomass in each area. We chose to use the output from the standard global algorithm to simplify the comparison of regions and of various years, to facilitate a comparison among all regions, and to avoid problems of defi ning boundaries of optically different regions. While this approach may introduce error into absolute estimates of productivity within a region, it provides a uniform basis to compute productivity throughout the Southern Ocean, as regional algorithms (some of which need more rigorous validation) are not available for all areas. RESULTS SPATIAL MEANS AND VARIABILITY Annual productivity of the Southern Ocean is highly variable but also quite low relative to other oceans, as has been suggested based on discrete measurements (e.g., Smith and Nelson, 1986; Nelson et al., 1996; Tremblay and Smith, 2007). Much of the region off the continental shelf is oligotrophic and is characterized by primary production rates of less than 50 g C m H110021 y H110021 (Figure 1). Regions of FIGURE 1. Mean (1998? 2006) modeled productivity of the Southern Ocean as derived from a vertically integrated productivity model. 23_Smith_pg309-318_Poles.indd 31123_Smith_pg309-318_Poles.indd 311 11/17/08 8:44:18 AM11/17/08 8:44:18 AM 312 SMITHSONIAN AT THE POLES / SMITH AND COMISO enhanced (threefold greater than the low-productivity off- shore areas) do occur on the continental shelf, with three areas being noteworthy: the Ross Sea, the Amundsen Sea, and Prydz Bay/East Antarctic shelf. Productivity in the Ross Sea is spatially extensive, but the greatest absolute productivity is in the Amundsen Sea region (150 g C m H110021 y H110021 at H1101173?S, 110?W). It is interesting that this particular region has never been sampled because of the diffi culty of gaining access by ships. Productivity in the more northern regions (near the location of the Antarctic Circumpolar Current (ACC) and its associated fronts, e.g., Abbott et al., 2000) is elevated in the Pacifi c sector (between 45? and 135?W) and south of New Zealand (between 155?W and 165?E), averaging H1101175 g C m H110021 y H110021 , and can be contrasted with the very low productivity waters of the South Atlantic and Indian Ocean sectors (Figure 1). Productivity of the South Atlantic is greater farther north than that in our selected study re- gion (58?S; Moore and Abbott, 2000; Smith and Comiso, 2008), and the region we analyzed is also largely south of the ACC (Moore and Abbott, 2000) and largely free of frontal enhancements. The Indian Ocean sector is among the windiest areas on Earth, and hence deep mixing would be expected to occur. Regardless, the annual productivity in the southern Indian Ocean and South Atlantic areas is less than 20 g C m H110021 y H110021 , among the lowest anywhere in the world?s oceans. Computed standard deviations for the entire South- ern Ocean suggest that while the absolute variations occur in the most productive continental shelf regions such as the Ross Sea, the relative spatial variations are actually greater elsewhere (Figure 2). For example, in the Ross Sea the standard deviation expressed as a percentage of the mean is only 2.8%, whereas in the southern Weddell Sea they range from 5.4% to 20%, suggesting the spatial vari- ability in that location is much greater. This likely is due to the impact of ice, which varies greatly in this location interannually (Smith and Comiso, 2008). The highest pro- ductivity occurs in areas of polynyas; in the Ross Sea the standard deviation is not as large because the location of the polynya is basically the same from one year to another. Variations in the Amundsen Sea, the location of the pro- ductivity maximum, are also less than in other regions, be- ing similar to those in the Ross Sea (H110111.5%? 3%). Varia- tions in the South Atlantic can be substantial (H1101110% near the location of the Weddell Sea polynya and Maud Rise) as well. Conversely, the elevated productivity region in the Pacifi c sector north of 62?S exhibits quite low variability (generally less than 1%). SEASONAL PRODUCTIVITY AND VARIABILITY ON MONTHLY SCALES The broad seasonal progression of productivity in some regions of the Antarctic is relatively well known. For example, in the Ross Sea a rapid increase in phytoplank- ton biomass and productivity occurs in spring, and a de- cline begins in mid-December to early January. Much of the summer is characterized by relatively low biomass and productivity (Smith et al., 2000, 2003). Productivity in the West Antarctic Peninsula region also is characterized by a similar pattern (Ducklow et al., 2006), although the mag- nitude of the productivity is far less (Smith and Comiso, 2008). December productivity in the Southern Ocean parallels the annual pattern, with the maximum produc- tivity occurring in the Amundsen and Ross seas and East Antarctic continental shelf (Figure 3). Clearly, the high- productivity areas are those of the continental shelf. Pro- ductivity north of 62?S is also higher in the Pacifi c sector. January productivity is characterized by increased rates and spatial extents in the Amundsen and Weddell seas, as well as at the tip of the Antarctic Peninsula, but by a decrease in the Ross Sea. February rates show a general decrease, with decreases being most noticeable in the East Antarctic region, the peninsula area, Amundsen Sea, and FIGURE 2. Standard deviation of the derived annual productivity values. Only those pixels where there were at least fi ve years of data (from 1998 to 2006) were included. Black regions are those with fewer than fi ve values; white areas have no data. 23_Smith_pg309-318_Poles.indd 31223_Smith_pg309-318_Poles.indd 312 11/17/08 8:44:29 AM11/17/08 8:44:29 AM SOUTHERN OCEAN PRIMARY PRODUCTIVITY 313 the Pacifi c sector north of 62?S. All sites show the general- ized maximum in late spring or early summer, followed by a decrease, although the timing of various sites varies. Variability on a monthly basis appears to be larger than on an annual basis (Figure 4). For example, relative variations in the Ross Sea are H110117% in all months, suggest- ing that the annual variations are somewhat dampened by the effects of long low-productivity periods. December variations are diffi cult to assess, as many locations have fewer than fi ve years of data and hence no standard devia- tion was calculated. However, variability in general seems to increase slightly in February, which may refl ect the rel- atively stochastic occurrence of storms (and hence deep mixing) during that period. DISCUSSION Primary productivity estimates in the Southern Ocean have been made for decades but have resulted in a biased picture of photosynthesis and growth. This is largely be- cause historically, estimates have been made in ice-free waters (e.g., Holm-Hansen et al., 1977; El-Sayed et al., 1983), whereas polynyas, which are known to be sites of intensive productivity (Tremblay and Smith, 2007), have rarely been sampled. Additionally, open water regions of low production have largely been ignored, and sam- pling has concentrated on the high-productivity locations thought to support local food webs. The richness of upper trophic levels that has been observed for over 100 years (e.g., Knox, 1994) was so marked that it was assumed that primary production must occur to support this abun- dance. However, we now recognize that productivity in the Southern Ocean is not great (Smith and Nelson, 1986), particularly on an annual basis, and the abundant higher trophic level standing stocks and extensive biogenic sedi- mentary deposits are forced by food web effi ciency, alter- nate food sources, and uncoupling of carbon with silica in biogeochemical cycles (Nelson et al., 1996). With the advent of satellite oceanography, large, syn- optic measurements of phytoplankton biomass became available. Such estimates in the Southern Ocean were far less common than in tropical and temperate waters, but they were very useful in showing the relationship of chlo- rophyll with ice distributions (Nelson et al., 1987; Sullivan et al., 1993), hydrographic features and fronts (Moore and Abbott, 2000), and depth (Comiso et al., 1993). In general, early satellite studies suggested that coastal zones and marginal ice zones were sites of large phytoplankton FIGURE 3. Mean monthly productivity of the Southern Ocean for the years 1998? 2006 for (a) December, (b) January, and (c) February. 23_Smith_pg309-318_Poles.indd 31323_Smith_pg309-318_Poles.indd 313 11/17/08 8:44:40 AM11/17/08 8:44:40 AM 314 SMITHSONIAN AT THE POLES / SMITH AND COMISO biomass accumulation (Sullivan et al., 1993; Arrigo and McLain, 1994). More refi ned treatments suggested that the Southern Ocean had a number of hot spots and short- lived increases in biomass (Moore et al., 1999) but, in large part, was extremely oligotrophic in nature. For many years it was uncertain why the Antarc- tic was so oligotrophic. Many considered that vertical mixing created low-irradiance conditions, superimposed on the seasonal aspects of ice distributions and solar angle, both which restricted irradiance penetration into the surface (e.g., Smith and Nelson, 1985; Mitchell and Holm- Hansen, 1991). Macronutrients such as nitrate and phosphate were always in excess, and it was sug- gested that micronutrients such as iron or vitamin B-12 might limit production (e.g., Hart, 1934). However, reli- able data on the concentrations of these micronutrients was lacking until the 1990s, when trace-metal clean mea- surements were made (e.g., Martin et al., 1990; Fitzwater et al., 2000). Iron concentrations were indeed found to be vanishingly small? in many cases less than 0.3 nM, even in coastal regions (Sedwick and DiTullio, 1997; Sedwick et al., 2000; Boyd and Abraham, 2001; Coale et al., 2003; de Baar et al., 2005). Furthermore, on the basis of laboratory studies and then fi eld work, under low-irradiance conditions, iron demands increase; hence, the interactive effects between iron and light exacerbated the limitation, and this interaction was suggested to be of paramount importance in deeper, offshore regions (Boyd and Abraham, 2001; Smith and Comiso, 2008). Recently, it has been found that vitamin B-12 can limit or colimit phytoplankton growth in the Ross Sea (Bertrund et al., 2007), but the large-scale colimitation for the en- tire Southern Ocean remains to be demonstrated. Other potential productivity-limiting factors have been addressed as well, such as grazing (Tagliabue and Arrigo, 2003) and temperature. However, herbivore biomass in- ventories are available only in selected regions and hence cannot be extrapolated over the entire Antarctic; further- more, the effects of temperature have been considered to be of secondary importance in limiting growth and photo- synthesis (Arrigo, 2007), although temperature may have a signifi cant role in controlling assemblage composition. It is useful to compare satellite means with other esti- mates that have been made, either via in situ measurements or numerical models. However, there are surprisingly few regions in the Southern Ocean that have adequate time series data to resolve the annual production signal; simi- larly, few regions have been the focus of intensive model- ing. One region that has received assessments from both detailed measurements and numerical modeling is the FIGURE 4. Standard deviations of the monthly productivity of the Southern Ocean for the years 1998? 2006 for (a) December, (b) Jan- uary, and (c) February. Black regions are those with fewer than fi ve values; white areas have no data. 23_Smith_pg309-318_Poles.indd 31423_Smith_pg309-318_Poles.indd 314 11/17/08 8:44:57 AM11/17/08 8:44:57 AM SOUTHERN OCEAN PRIMARY PRODUCTIVITY 315 Ross Sea. Tremblay and Smith (2007, table 2) used the nutrient climatology compiled by Smith et al. (2003) and estimated the productivity by month and by year. The an- nual productivity based on nitrogen uptake was 155 g C m H110021 y H110021 , remarkably similar to the value estimated from our satellite model (Table 1). Smith and Gordon (1997) used measurements taken during November, along with other estimates, and calculated production to be 134 g C m H110021 y H110021 . Arrigo et al. (1998), using a numerical model, estimated productivity to be H11011160 g C m H110021 y H110021 . The simi- larity between all of these estimates, either direct or indi- rect, and ours derived from satellite estimates and a verti- cally integrated production model is striking and gives us confi dence that our procedure accurately assesses the pro- duction, despite the suggestion that chlorophyll retrievals from space in this area may contain signifi cant errors (Ar- rigo et al., 1998). As the Ross Sea is the Antarctic?s most spatial extensive phytoplankton bloom, the mean annual productivity is also near the maximum for the Antarctic. Our results suggest that the productivity of the Amundsen Sea may be slightly greater. The region is the site of a num- ber of spring polynyas, and the optical properties of the water are likely similar to those in the Ross Sea. However, currently there are very limited in situ measurements avail- able to confi rm this substantial productivity. It has been suggested that the high productivity of the Ross Sea is derived from substantial vertical stratifi cation, early removal of ice, and adequate macro- and micronu- trients for much of the growing season (Smith and Asper, 2001; Smith et al., 2003, 2006), coupled with limited grazing (Tagliabue and Arrigo, 2003). It has also been shown that during some summers a large ?secondary? bloom occurs (Peloquin and Smith, 2007) and that these blooms occur approximately every three years. Peloquin and Smith (2007) suggested that summer iron limitation is occasionally reduced or eliminated by the intrusion of Modifi ed Circumpolar Deep Water onto the continental shelf by oceanographic processes. Such a process would contribute greatly to the increased February variability we observed at some locations. A similar pattern of oceanic circulation has been suggested for the Prydz Bay/East Ant- arctica region as well (Smith et al., 1984), and it would be interesting to know if a similar infl uence of currents is responsible for the high productivity we observed in the Amundsen Sea. TEMPORAL PATTERNS OF PRODUCTIVITY IN THE SOUTHERN OCEAN The data that are used to derive the mean productivity shown in Figure 1 have also been analyzed for temporal trends (Figure 5). Mean Antarctic productivity for the past decade has shown a signifi cant increase; furthermore, this increase is driven by changes that are largely confi ned to January and February (Smith and Comiso, 2008). Mod- els have suggested that the productivity of the Southern Ocean will increase under atmospheric temperature in- creases driven by CO 2 loading (Sarmiento and Le Qu?r?, 1996; Sarmiento et al., 1998; Behrenfeld et al., 2006). The change will result from increased ice melting, which, in turn, should increase stratifi cation, rather than a direct tem- perature effect. An increase in stratifi cation would increase TABLE 1. Summary of annual productivity estimates and method of computation for various portions of the Southern Ocean. ? Southern? means the assessment was confi ned to regions south of 75?S. Region Annual productivity (g C m H110022 y H110021 ) Method of estimation Reference Ross Sea 112 biomass accumulation Nelson et al. (1996) Ross Sea (southern) 190 biomass accumulation Smith and Gordon (1997) Southern Ocean 95.4? 208 bio-optical model Behrenfeld and Falkowski (1997a) Southern Ocean 105 bio-optical model Arrigo et al. (1998) Ross Sea 57.6 H11006 22.8 nutrient defi cits Sweeney et al. (2000) Southern Ocean 62.4 bio-optical model Moore and Abbott (2000) Ross Sea 151 H11006 21 bio-optical model Arrigo and van Dijken (2003) Ross Sea (southern) 145 numerical model Arrigo et al. (2003) Ross Sea 84? 218 nutrient defi cits Smith et al. (2006) Ross Sea (southern) 153 nutrient defi cits Tremblay and Smith (2007) Ross Sea (southern) 54? 65 bio-optical model Smith and Comiso (2008) Southern Ocean 20? 150 bio-optical model this study 23_Smith_pg309-318_Poles.indd 31523_Smith_pg309-318_Poles.indd 315 11/17/08 8:45:16 AM11/17/08 8:45:16 AM 316 SMITHSONIAN AT THE POLES / SMITH AND COMISO the net irradiance environment available to phytoplank- ton and also decrease the magnitude of the iron-irradiance interaction, resulting in a decreased iron demand. Should iron inputs and concentrations remain the same, then the increased productivity would result in large-scale increases in phytoplankton growth and productivity. The observed change in the productivity estimated from satellites are not necessarily indicative of the changes predicted by the mod- els and may refl ect shorter-term trends that have altered current patterns, atmospheric inputs of iron, or other fac- tors. It should also be noted that models do not include any colimitation effects of vitamin B-12, and if this effect were signifi cant throughout the Southern Ocean, then the increase in productivity would be smaller than predicted. Regardless, the observed increase in annual productivity was unexpected, and the data analysis should be continued (using the same methods) as far into the future as possible to confi rm this pattern. Smith and Comiso (2008) also attempted to ascertain if the satellite data could be used to detect changes in pro- ductivity on a regional scale. Given that certain regions are having signifi cant alterations in ice concentrations (e.g., the West Antarctic Peninsula? Amundsen/ Bellingshausen Sea sector has had a H110227% decrease per decade in ice concen- tration, while the Ross Sea sector has had an increase of H110225%; Kwok and Comiso, 2002), it might be expected that changes in productivity are accompanying these changes. However, the temporal variability in the estimates of pro- ductivity of these areas was too great to allow for any trends to be determined, so at this time, it is impossible to determine if changes in higher trophic levels are occurring because of food web effects (via energetics) or by habitat modifi cation (e.g., loss of reproductive sites and decreases in reproductive success). CONCLUSIONS Tremendous advances have been made in our under- standing of primary productivity in the Southern Ocean in the past 50 years. We have moved from an era of observa- tional science into one that combines observations and ex- periments with large-scale assessments using data derived from multiple satellites and modeling using the same data. We recognize that the earlier assessments of productivity were biased by sampling and the nature of Antarctic pro- ductivity, and using unbiased techniques such as satellite data combined with robust models provides a means by which the temporal and spatial trends in phytoplankton production can be assessed. These methods have clearly demonstrated that the Southern Ocean as a whole is an oligotrophic area, with enhanced productivity on the continental shelves. Yet the shelf productivity is far from evenly distributed, and it is likely that oceanographic in- fl uences may play a large role in setting the maximum lim- its to production in the Southern Ocean. It is suggested that the productivity of the entire South- ern Ocean has increased signifi cantly in the past decade, although the causes for such an increase remain obscure. Such changes have been predicted by numerical models, but it is far from certain that the observed changes are in fact related to climate change in the Antarctic. The short- term record also makes it diffi cult to interpret what the trend really means, especially in light of the possible ef- fect of some climate modes like the Southern Hemisphere Annular Mode (Kwok and Comiso, 2002; Gordon et al., 2007). Only through extended analyses can such trends be confi rmed and the causes for these changes ascertained. While increases in productivity of the magnitude shown may not induce major shifts in the ecology and biogeo- chemistry of the region, such changes, if they continue, may result in subtle and unpredicted impacts on the foods webs of the Antarctic ecosystem as well as changes in el- emental dynamics. Knowledge of the environmental reg- ulation of these changes in productivity is critical to the understanding of the ecology of the entire Southern Ocean FIGURE 5. The temporal pattern of productivity for the entire Southern Ocean as derived from satellite data and a productivity model (from Smith and Comiso, 2008). The trend is derived from a linear regression of all points and is highly signifi cant. 23_Smith_pg309-318_Poles.indd 31623_Smith_pg309-318_Poles.indd 316 11/17/08 8:45:16 AM11/17/08 8:45:16 AM SOUTHERN OCEAN PRIMARY PRODUCTIVITY 317 and will provide insights into the potential changes that will undoubtedly occur in the coming years. ACKNOWLEDGMENTS The expert computer skills of Larry Stock of STX are gratefully acknowledged. This work was funded partially by National Sciences Foundation grants OPP-0087401 and OPP-0337247 (to WOS) and by the NASA Cryo- spheric Science Program (to JCC). This is VIMS contribu- tion number 2955. LITERATURE CITED Abbott, M. R., J. G. Richman, R. M. Letelier, and J. S. Bartlett. 2000. The Spring Bloom in the Antarctic Polar Frontal Zone as Observed from a Mesoscale Array of Bio-optical Sensors. 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Robinson, G. C. Geldman, S. W. Bailey, J. Gales, P. J. Werdell, M. Wang, R. Frouin, R. P. Stumpf, R. A. Arnone, R. W. Gould Jr., P. M. Martino- lich, V. Ramsibrahmanakul, J. E. O?Reilly, and J. A. Yoder. 2003. Algorithm Updates for the Fourth SeaWiFS Data Reprocessing. NASA Technical Memorandum NASA/ TM-2003-206892. NASA Goddard Space Flight Center, Greenbelt, Md. Peloquin, J. A., and W. O. Smith Jr. 2007. Phytoplankton blooms in the Ross Sea, Antarctica: Interannual Variability in Magnitude, Tempo- ral Patterns, and Composition. Journal of Geophysical Research, 112: C08013, doi: 10.1029/ 2006JC003816. Sarmiento, J. L., T. M. C. Hughes, R. J. Stouffer, and S. Manabe.1998. Simulated Response of the Ocean Carbon Cycle to Anthropogenic Climate Warming. Nature, 393: 245? 249. Sarmiento, J. L., and C. Le Qu?r?. 1996. Oceanic Carbon Dioxide Up- take in a Model of Century-Scale Global Warming. Science, 274: 1346? 1350. Sedwick, P. N., and G. R. DiTullio. 1997. Regulation of Algal Blooms in Antarctic Shelf Waters by the Release of Iron from Melting Sea Ice. Geophysical Research Letters, 24: 2515? 2518. Sedwick, P. N., G. R. DiTullio, and D. J. Mackey. 2000. Iron and Man- ganese in the Ross Sea, Antarctica: Seasonal Iron Limitation in Antarctic Shelf Waters. Journal of Geophysical Research, 105: 11321? 11336. Smith, N. R., D. Zhaoqian, K. R. Kerry, and S. Wright. 1984. Water Masses and Circulation in the Region of Prydz Bay, Antarctica. Deep-Sea Research, Part A, 31: 1121? 1137. Smith, W. O., Jr., and V. A. Asper. 2001. The Infl uence of Phytoplankton Assemblage Composition on Biogeochemical Characteristics and Cycles in the Southern Ross Sea, Antarctica. Deep-Sea Research, Part I, 48:137? 161. Smith, W. O., Jr., and J. C. Comiso. 2008. Infl uence of Sea Ice on Primary Production in the Southern Ocean: A Satellite Perspective. Journal of Geophysical Research, 113: C05S93, doi: 10.1029/ 2007JC004251. Smith, W. O., Jr., M. S. Dinniman, J. M. Klinck, and E. Hofmann. 2003. Biogeochemical Climatologies in the Ross Sea, Antarctica: Seasonal Patterns of Nutrients and Biomass. Deep-Sea Research, Part II, 47: 3083? 3101. Smith, W. O., Jr., and L. I. Gordon. 1997. Hyperproductivity of the Ross Sea (Antarctica) Polynya during Austral Spring, Geophysical Re- search Letters, 24: 233? 236. Smith, W. O., Jr., J. Marra, M. R. Hiscock, and R. T. Barber. 2000. The Seasonal Cycle of Phytoplankton Biomass and Primary Productiv- ity in the Ross Sea, Antarctica. Deep-Sea Research. Part II, 47: 3119? 3140. Smith, W. O., Jr., and D. M. Nelson. 1985. Phytoplankton Bloom Pro- duced by a Receding Ice Edge in the Ross Sea: Spatial Coherence with the Density Field. Science, 227: 163? 166. ???. 1986. The Importance of Ice-Edge Blooms in the Southern Ocean. BioScience, 36: 251? 257. Smith, W. O., Jr., A. R. Shields, J. A. Peloquin, G. Catalano, S. Tozzi, M. S. Dinniman, and V. A. Asper. 2006. Biogeochemical Budgets in the Ross Sea: Variations among Years. Deep-Sea Research, Part II, 53: 815? 833. Sullivan, C. W., K. R. Arrigo, C. R. McClain, J. C. Comiso, and J. Fire- stone. 1993. Distributions of Phytoplankton Blooms in the South- ern Ocean. Science, 262: 1832? 1837. Sunda, W. G., and S. A. Huntsman. 1997. Interrelated Infl uence of Iron, Light and Cell Size on Marine Phytoplankton Growth. Nature, 390: 389? 392. Sweeney, C., D. A. Hansell, C. A. Carlson, L. A. Codispoti, L. I. Gordon, J. Marra, F. J. Millero, W. O. Smith Jr., and T. Takahashi. 2000. Biogeochemical Regimes, Net Community Production and Carbon Export in the Ross Sea, Antarctica. Deep-Sea Research, Part II, 47: 3369? 3394. Tagliabue, A., and K. R. Arrigo. 2003. Anomalously Low Zooplankton Abundance in the Ross Sea: An Alternative Explanation. Limnol- ogy and Oceanography, 48: 686? 699. Tremblay, J.-E., and W. O. Smith Jr. 2007. ?Phytoplankton Processes in Polynyas.? In Polynyas: Windows to the World?s Oceans, ed. W. O. Smith Jr. and D. G. Barber, pp. 239? 270. Amsterdam: Elsevier. Vaughan, D. G., G. J. Marshall, W. M. Connolley, C. Parkinson, R. Mul- vaney, D. A. Hodgson, J.C. King, C. J. Pudsey, and J. Turner. 2003. Recent Rapid Regional Climate Warming on the Antarctic Penin- sula. Climatic Change, 60: 243? 274. 23_Smith_pg309-318_Poles.indd 31823_Smith_pg309-318_Poles.indd 318 11/17/08 8:45:18 AM11/17/08 8:45:18 AM ABSTRACT. Chromophoric dissolved organic matter (CDOM) is ubiquitous in the oceans, where it is an important elemental reservoir, a key photoreactant, and a sun- screen for ultraviolet (UV) radiation. Chromophoric dissolved organic matter is generally the main attenuator of UV radiation in the water column, and it affects the remote sens- ing of chlorophyll a (chl a) such that corrections for CDOM need to be incorporated into remote sensing algorithms. Despite its signifi cance, relatively few CDOM measurements have been made in the open ocean, especially in polar regions. In this paper, we show that CDOM spectral absorption coeffi cients (a ? ) are relatively low in highly productive Antarctic waters, ranging from approximately 0.18 to 0.30 m H110021 at 300 nm and 0.014 to 0.054 m H110021 at 443 nm. These values are low compared to coastal waters, but they are higher (by approximately a factor of two to three) than a ? in oligotrophic waters at low latitudes, supporting the supposition of a poleward increase in a CDOM in the open ocean. Chromophoric dissolved organic matter a ? and spectral slopes did not increase during the early development of a bloom of the colonial haptophyte Phaeocystis antarctica in the Ross Sea, Antarctica, even though chl a concentrations increased more than one- hundred-fold. Our results suggest that Antarctic CDOM in the Ross Sea is not coupled directly to algal production of organic matter in the photic zone during the early bloom but is rather produced in the photic zone at a later time or elsewhere in the water column, possibly from organic-rich sea ice or the microbial degradation of algal-derived dissolved organic matter exported out of the photic zone. Spectral a ? at 325 nm for surface waters in the Southern Ocean and Ross Sea were remarkably similar to values reported for deep water from the North Atlantic by Nelson et al. in 2007. This similarity may not be a coin- cidence and may indicate long-range transport to the North Atlantic of CDOM produced in the Antarctic via Antarctic Intermediate and Bottom Water. INTRODUCTION Quantifying temporal and spatial variations in chromophoric dissolved or- ganic matter (CDOM) absorption is important to understanding the biogeo- chemistry of natural waters because CDOM plays a signifi cant role in deter- mining the underwater light fi eld. Chromophoric dissolved organic matter is the fraction of dissolved organic matter (DOM) that absorbs solar radiation in natural waters, including radiation in the UVB (280 to 320 nm), the UVA (320 to 400 nm), and the visible portion (400 to 700 nm) of the solar spectrum. For most natural waters, CDOM is the primary constituent that attenuates actinic David J. Kieber, Department of Chemistry, State University of New York College of Environmental Science and Forestry, Syracuse, NY 13210, USA. Dierdre A. Toole, Department of Marine Chem- istry and Geochemistry, Woods Hole Oceano- graphic Institution, Woods Hole, MA 02543, USA. Ronald P. Kiene, Department of Marine Sci- ences, University of South Alabama, Mobile, AL 36688, USA. Corresponding author: D. Kieber (djkieber@esf.edu). Accepted 28 May 2008. Chromophoric Dissolved Organic Matter Cycling during a Ross Sea Phaeocystis antarctica Bloom David J. Kieber, Dierdre A. Toole, and Ronald P. Kiene 24_Kieber_pg319-334_Poles.indd 31924_Kieber_pg319-334_Poles.indd 319 11/18/08 9:15:26 AM11/18/08 9:15:26 AM 320 SMITHSONIAN AT THE POLES / KIEBER, TOOLE, AND KIENE UVB and UVA radiation in the water column. Conse- quently, CDOM acts as a ?sunscreen,? providing protec- tion from short wavelengths of solar radiation that can be damaging to aquatic organisms (Hebling and Zagarese, 2003). Chromophoric dissolved organic matter absorp- tion is often also quite high in the blue spectral region, particularly in the subtropics and poles, accounting for a quantitatively signifi cant percentage of total nonwater absorption (Siegel et al., 2002), complicating remote sens- ing of phytoplankton pigment concentrations and primary productivity (Carder et al., 1989; Nelson et al., 1998). In lakes and coastal areas with high riverine discharge, CDOM absorption in the blue region can be so large as to restrict phytoplankton absorption of light, thereby placing limits on primary productivity (e.g., Vodacek et al., 1997; Del Vecchio and Blough, 2004). Since CDOM absorption affects the satellite-retrieved phytoplankton absorption signal in most oceanic waters, especially coastal waters, CDOM absorption must be accounted for when using re- motely sensed data (Blough and Del Vecchio, 2002). This necessitates a thorough understanding of the characteris- tics of CDOM and factors controlling its distribution in different areas of the ocean. Sources of CDOM in the oceans are varied and, in many cases, poorly described. Potential sources include terrestrial input of decaying plant organic matter, autochthonous pro- duction by algae, photochemical or bacterial processing of DOM, and release from sediments (e.g., Blough et al., 1993; Opsahl and Benner, 1997; Nelson et al., 1998; Nelson and Siegel, 2002; Rochelle-Newall and Fisher, 2002a, 2002b; Chen et al., 2004; Nelson et al., 2004). The removal of CDOM in the oceans is dominated by its photochemical bleaching, but microbial processing is also important al- though poorly understood (Blough et al., 1993; Vodacek et al., 1997; Del Vecchio and Blough, 2002; Chen et al., 2004; Nelson et al., 2004; V?h?talo and Wetzel, 2004). Absorption of sunlight by CDOM can affect an aquatic ecosystem both directly and indirectly (Schindler and Cur- tis, 1997). The bleaching of CDOM absorption and fl uo- rescence properties as a result of sunlight absorption lessens the biological shielding effect of CDOM in surface waters. Chromophoric dissolved organic matter photobleaching may also produce a variety of reactive oxygen species (ROS), such as the hydroxyl radical, hydrogen peroxide, and superoxide (for review, see Kieber et al., 2003). Gen- eration of these compounds provides a positive feedback to CDOM removal by causing further destruction of CDOM via reaction with these ROS. Although CDOM has been intensively studied to understand its chemical and physical properties, previ- ous research has primarily focused on CDOM cycling in temperate and subtropical waters. Very little is known re- garding temporal and spatial distributions of CDOM in high-latitude marine waters. Studies in the Bering Strait? Chukchi Sea region and the Greenland Sea showed that absorption coeffi cients (a ? ) and spectral slopes (S) derived from CDOM absorption spectra at coastal stations were infl uenced by terrestrial inputs and comparable to a ? and S values in temperate and tropical coastal sites (Stedmon and Markager, 2001; Ferenac, 2006). As the distance from the coastline in the Bering Strait and Chukchi Sea proper increased, there was a large decrease in a ? from 1.5 to 0.2 m H110021 at 350 nm. These a ? were mostly higher than a 350 observed in the open ocean at lower latitudes (Ferenac, 2006). However, corresponding spectral slopes were signifi cantly lower (H113490.014 nm H110021 ) than open oceanic values in the Sargasso Sea and elsewhere (H113500.02 nm H110021 ; Blough and Del Vecchio, 2002), suggesting that the types of CDOM present in these contrasting environments were different, possibly owing to a residual terrestrial signal and lower degree of photobleaching in the Arctic samples. As with Arctic waters, there is very little known regard- ing CDOM in Antarctic waters. To our knowledge, there are only three published reports investigating CDOM dis- tributions in the Ross Sea and along the Antarctic Peninsula (Sarpal et al., 1995; Patterson, 2000; Kieber et al., 2007). A key fi nding of the Sarpal et al. study was that Antarctic wa- ters were quite transparent in the UV (a ? H11021 0.4 m H110021 at ? H11350 290 nm), even at coastal stations during a bloom of Cryp- tomonas sp. Patterson (2000) examined CDOM in several transects perpendicular to the Antarctic Peninsula coastline and found that CDOM absorption coeffi cients were low in the austral summer, with a 305 (H11006SD) H11005 0.28 (H110060.09) m H110021 and a 340 H11005 0.13 (H110060.06) m H110021 . Similarly low a ? were ob- served by Kieber et al. (2007) in the upper water column of the Ross Sea (a 300 H11011 0.32 m H110021 and a 350 H11011 0.15 m H110021 ) during the end of a Phaeocystis antarctica bloom when diatoms were also blooming (chl a H11011 3.8 H9262g L H110021 ). Polar regions are unique in many ways that may infl u- ence CDOM optical properties. Sea ice may provide a rich source of ice-derived CDOM to melt water (Scully and Miller, 2000; Xie and Gosselin, 2005). Additionally, polar blooms can decay without signifi cant losses to zooplank- ton, especially in the spring when zooplankton grazing can be minimal (Caron et al., 2000; Overland and Stabeno, 2004; Rose and Caron, 2007). In the Ross Sea, soon after the opening of the Ross Sea polynya in the austral spring, a Phaeocystis antarctica? dominated bloom regularly occurs (Smith et al., 2000), especially in waters away from the ice edge where iron 24_Kieber_pg319-334_Poles.indd 32024_Kieber_pg319-334_Poles.indd 320 11/18/08 9:15:26 AM11/18/08 9:15:26 AM CHROMOPHORIC DISSOLVED ORGANIC MATTER CYCLING 321 concentrations are lower and the water column is more deeply mixed. As is typical of polar regions, the massive phytoplankton production that is observed during the early stages of this bloom (in the early to mid austral spring) is not accompanied by signifi cant micro- or macrozooplank- ton grazing (Caron et al., 2000). The spring bloom also appears to be a period of low bacterial abundance and ac- tivity (Ducklow et al., 2001), although bacteria do bloom during the later stages of the algal bloom, in early to mid summer, coinciding with an increase in dissolved and par- ticulate organic carbon (Carlson et al., 2000). Depending on physiological and hydrodynamic fac- tors, this ecological decoupling between primary produc- tivity and both microbial productivity and grazing may cause the phytoplankton to sink out of the photic zone (DiTullio et al., 2000; Becquevort and Smith, 2001; Over- land and Stabeno, 2004). This may lead to a signifi cant fl ux of organic matter to the ocean fl oor and to deepwater CDOM production. Export of the phytoplankton out of the photic zone may also lead to a decoupling of bloom dynamics and CDOM cycling. Here we report on spatial and temporal patterns in CDOM spectra observed during the early stages of the 2005 austral spring Phaeocystis antarctica bloom in the seasonal Ross Sea polynya. Our results show that CDOM changed very little during a period when chl a concentrations in- creased more than one-hundred-fold. Implications of this fi nding for CDOM cycling in the Ross Sea are discussed. METHODS ROSS SEA SITE DESCRIPTION AND SAMPLING A fi eld campaign was conducted aboard the R/V Nathaniel B. Palmer in the seasonal ice-free polynya in the Ross Sea, Antarctica, from 8 November to 30 November 2005 (Figure 1). During this time, the surface seawater tem- perature was approximately ? 1.8?C and the phytoplank- ton assemblage was dominated by colonial Phaeocystis antarctica. Three main hydrographic stations were occu- pied within the Ross Sea polynya for approximately seven days each (i.e., R10, R13, R14). Seawater samples were collected from early morning hydrocasts (0400-0700 local time) directly from Niskin bottles attached to a conductiv- ity, temperature, and depth (CTD) rosette. Vertical profi les of a suite of routine measurements were obtained at each station including downwelling irradiance (and coupled in- cident surface irradiance), chl a, and CDOM absorption spectra. Additionally, a profi le of dissolved organic carbon (DOC) was collected at one station (14F). In addition to studying CDOM cycling in the Ross Sea, we also collected and analyzed surface water CDOM samples during our North-South transit from New Zealand through the Southern Ocean to the Ross Sea (28 October? 7 November, 2005). Transect sampling for CDOM was con- ducted from H1101149 to H1101175?S. Details of the cruise track, sam- pling protocols, analyses conducted and meteoro logical and sea ice conditions for the southbound transect are presented in Kiene et al. (2007). The transect encompassed open waters of the Southern Ocean, the northern sea-ice melt zone, and ice-covered areas near the northern Ross Sea. Chlorophyll a was determined fl uorometrically by em- ploying the acidifi cation method described by Strickland and Parsons (1968). Briefl y, 50? 250 mL of seawater was fi l- tered onto a 25-mm GF/C glass fi ber fi lter (Whatman Inc., Floram Park, New Jersey) with low vacuum. Filters were placed in 5 mL of 90% HPLC-grade acetone and extracted for 24 h at ? 20?C. Chlorophyll fl uorescence in the acetone extracts was quantifi ed with a Turner Designs 10-AU fl uo- rometer (Sunnyvale, Calafornia) before and after 80-H9262L ad- dition of 10% HCl. Dissolved organic carbon samples were FIGURE 1. Locations of transect sampling stations south of New Zealand to the Ross Sea (open circles) and the main hydrographic stations that were occupied in the Ross Sea (solid black circles) dur- ing the NBP05-08 cruise. Latitude and longitude information for specifi c hydrographic stations (e.g., R10A and R10D) are given in fi gure captions 4, 5, 8 and 9. 24_Kieber_pg319-334_Poles.indd 32124_Kieber_pg319-334_Poles.indd 321 11/18/08 9:15:27 AM11/18/08 9:15:27 AM 322 SMITHSONIAN AT THE POLES / KIEBER, TOOLE, AND KIENE collected and concentrations were determined by employ- ing the techniques outlined in Qian and Mopper (1996). For nutrient and CDOM samples, seawater was col- lected directly from Niskin bottles by gravity fi ltration through a 20-H9262m Nitex mesh (held in a 47-mm-diameter polycarbonate (PC) fi lter holder) followed by a precleaned 0.2-H9262m AS 75 Polycap fi lter capsule (nylon membrane with a glass microfi ber prefi lter enclosed in a polypropylene housing). Silicone tubing was used to attach the PC fi lter to the Niskin bottle spigot and the Polycap to the PC fi lter outlet. Nutrient samples were collected into 50-mL poly- propylene centrifuge tubes, while samples for CDOM were fi ltered into precleaned 80-mL Qorpak bottles sealed with Tefl on-lined caps (see Toole et al., 2003, for details regard- ing sample fi ltration and glassware preparation). Nutrient samples were stored frozen at H1100280?C until shipboard anal- ysis by standard fl ow-injection techniques. CDOM ABSORPTION SPECTRA Absorbance spectra were determined with 0.2-H9262m fi l- tered seawater samples that were warmed to room tem- perature in the dark immediately after they were collected and then analyzed soon thereafter (generally within 24 h). Absorbance spectra were determined in a 100-cm path length, Type II liquid capillary waveguide (World Preci- sion Instruments) attached to an Ocean Optics model SD2000 dual-channel fi ber-optic spectrophotometer and a Micropack DH 2000 UV-visible light source. Prior to analysis, a sample or blank was fi rst deaerated with ul- trahigh purity He in an 80-mL Qorpak bottle and then pulled through the capillary cell slowly with a Rainin Rab- bit peristaltic pump. Deaeration with He eliminated bub- ble formation in the sample cell, while increasing the pH slightly from 8.0 to 8.3. This pH change is not expected to affect absorption spectra, although this was not explicitly tested. The reference solution consisted of 0.2-H9262m-fi ltered 0.7-M NaCl prepared from precombusted (600?C, 24 h) high-purity sodium chloride (99.8%, Baker Analyzed) and dissolved in high-purity (18.2 MH9024 cm) laboratory water obtained from a Millipore Milli-Q ultrapure water system (Millipore Corp., Billerica, Massachusetts). The sodium chloride solution was used to approximately match the ionic strength of the Ross Sea seawater samples and mini- mize spectral offsets due to refractive index effects. Even though a sodium chloride solution was used as a reference, sample absorbance spectra still exhibited small, variable baseline offsets (H110110.005 AU). This was corrected for by adjusting the absorbance (A ? ) to zero between 650 and 675 nm where the sample absorbance was assumed to be zero. The capillary cell was fl ushed after every fourth or fi fth seawater sample with deaerated Milli-Q water and high-purity, distilled-in-glass-grade methanol (Burdick and Jackson, Muskegon, Michigan). Corrected absorbance values were used to calculate absorption coeffi cients: a ? H11005 2.303A ? / l, (1) where l is the path length of the capillary cell. Each sample was analyzed in triplicate, resulting in H113492% relative stan- dard deviation (RSD) in spectral absorbance values H11349400 nm. On the basis of three times the standard deviation of the sodium chloride reference, the limit of detection for measured absorption coeffi cients was approximately 0.002 m H110021 . To characterize sample absorption, spectra were fi t to the following exponential form (e.g., Twardowski et al., 2004): a ? H11005 a ? o e H11002S(?? ?o) , (2) where a ? o is the absorption coeffi cient at the reference wavelength, ? o , and S is the spectral slope (nm H110021 ). Data were fi t to a single exponential equation using SigmaPlot (SPSS Inc.). Slope coeffi cients were evaluated from 275 to 295 nm (S 275?295 ; ? o H11005 285 nm) and from 350 to 400 nm (S 350?400 ; ? o H11005 375 nm). These wavelength ranges were se- lected for study of spectral slopes instead of using broader wavelength ranges (e.g., 290? 700 nm) because the former can be measured with high precision and better refl ects biogeochemical changes in CDOM in the water column (Helms et al., 2008). OPTICAL PROFILES Vertical profi les of spectral downwelling irradiance (E d (z,?)) and upwelling radiance (L u (z,?)), as well as the time course of surface irradiance (E d (0 H11001 ,?, t)), were de- termined separately for ultraviolet and visible wavebands. Ultraviolet wavelengths were sampled using a Biospherical Instruments, Inc. (BSI, San Diego, California) PUV-2500 Profi ling Ultraviolet Radiometer coupled with a continu- ously sampling, deck-mounted, cosine-corrected GUV- 2511 Ground-based Ultraviolet Radiometer. Both sensors had a sampling rate of approximately 6 Hz and monitored seven channels centered at 305, 313, 320, 340, 380, and 395 nm, as well as integrated photosynthetically active radiation (PAR). Each channel had an approximate band- width of 10 nm, except for the 305-nm channel, whose bandwidth was determined by the atmospheric ozone cut- off and the PAR channel, which monitored the irradiance 24_Kieber_pg319-334_Poles.indd 32224_Kieber_pg319-334_Poles.indd 322 11/18/08 9:15:28 AM11/18/08 9:15:28 AM CHROMOPHORIC DISSOLVED ORGANIC MATTER CYCLING 323 from 400 to 700 nm. The irradiance at several visible wavelength channels, centered at 412, 443, 490, 510, 555, and 665 nm, as well as PAR, were determined with a BSI PRR-600 Profi ling Refl ectance Radiometer coupled to a radiometrically matched surface reference sensor (PRR- 610). Similarly, both PRR sensors had a sampling rate of approximately 6 Hz and an approximate bandwidth of 10 nm. The PUV-2500 was deployed multiple times a day (generally, three to fi ve profi les per day) in free-fall mode, allowing it to sample at a distance of over 10 m from the ship, minimizing effects from ship shadow and instrument tilt (Waters et al., 1990). The PRR-600 was deployed sev- eral times per station in a metal lowering frame via the starboard side winch. Prior to each cast, the ship was ori- ented relative to the sun to minimize ship shadow. The GUV-2511 and PRR-610 were mounted to the deck, and care was taken to avoid shadows or refl ected light associ- ated with the ship?s superstructure. To reduce instrument variability due to atmospheric temperature fl uctuations, the GUV-2511 was equipped with active internal heating. All calibrations utilized coeffi cients provided by BSI, and the data were processed using standard procedures. Spectral downwelling attenuation coeffi cients (K d (?), m H110021 ) were derived from each PUV-2500 and PRR-600 profi le as the slope of log-transformed E d (z, ?) versus depth. On the basis of water clarity, the depth interval for this calculation varied from H11021 10 m for shorter UV wavelengths up to 20-30 m for blue wavelengths of solar radiation. Daily mean K d (?) were derived as the average of individual K d (?) coeffi cients determined from each PUV and PRR profi le, and station means were determined by averaging the daily K d (?) coeffi cients. RESULTS SAMPLE STORAGE AND SPECTRAL COMPARISON A storage test was conducted with seawater collected on 29 October 2005 (52?59.90H11032S, 175?7.76H11032E) during the transect from New Zealand to the Ross Sea. The sample was obtained from the ship?s underway pump system that had an intake depth at approximately 4 m. Seawater was fi ltered directly from the pump line through a 0.2-?m AS 75 Polycap fi lter capsule into an 80-mL Qorpak bottle and stored in the dark at room temperature. When this sample was analyzed multiple times over a period of two weeks (n H11005 8), no change was observed in its absorp- tion spectrum with respect to spectral absorption coef- fi cients or spectral slopes. For example, a 300 varied over a very narrow range from 0.196 to 0.202 m H110021 with a mean value of 0.197 m H110021 and 3.1% RSD. This RSD is only slightly larger than the RSD for replicate analysis of the same sample when done sequentially (H110112%). Likewise, the spectral slope from 275 to 295 nm (S 275-295 ) showed very little variation over time, ranging from 0.0337 to 0.0377 nm H110021 (0.0358 nm H110021 mean and 3.4% RSD). The S 350-400 varied somewhat more (11.8% RSD) due to the lower a ? values, ranging from 0.0075 to 0.0111 nm H110021 , with a mean value of 0.009 nm H110021 . A storage study with two other samples that were collected along the transect showed similar results, with very little variability ob- served in either a ? or S beyond the precision of sequential spectral measurements. In addition to the storage study, absorption spectra obtained with the capillary waveguide system were com- pared to those obtained with the commonly used Perkin Elmer Lamda 18 dual-beam, grating monochromator spectrophotometer. There was no statistical difference in the spectral absorption coeffi cients obtained by these two spectrophotometers, except for considerably more noise in a ? obtained with the Perkin Elmer spectrophotometer (Figure 2). The increased spectral noise seen with the Beckman spectrophotometer was expected due to the rela- tively short path length (0.10 m) and the extremely low absorbance values in the Ross Sea seawater samples (e.g., H110110.009 AU at 400 nm), which were close to the detection limit of this instrument. In contrast, a ? spectra obtained FIGURE 2. Comparison of spectral absorption coeffi cients deter- mined with a Perkin-Elmer Lambda 18 dual-beam spectrophoto- meter (circles) and a capillary waveguide spectrophotometer (solid line). A 10-cm cylindrical quartz cell was used to obtain spectral absorption coeffi cients with the Perkin-Elmer spectrophotometer. Absorbance spectra were referenced against a 0.7 M NaCl solution. 24_Kieber_pg319-334_Poles.indd 32324_Kieber_pg319-334_Poles.indd 323 11/18/08 9:15:29 AM11/18/08 9:15:29 AM 324 SMITHSONIAN AT THE POLES / KIEBER, TOOLE, AND KIENE with the capillary waveguide were much smoother owing to the much higher absorbance values due to the much longer path length (1 m) and the ability to spectrally aver- age over several scans. TRANSECT AND ROSS SEA CDOM ABSORPTION SPECTRA Absorption coeffi cients obtained during our transect from New Zealand to the Ross Sea were generally low, as exemplifi ed by a 300 (Figure 3). Values for a 300 ranged from 0.178 to 0.264 m H110021 , with an average value of 0.209 m H110021 . Interestingly, absorption coeffi cients at 300 nm were consistently higher in samples collected with signifi cant ice cover (avg H11006 SD: 0.231 H11006 0.014 m H110021 ) compared to a 300 values collected in the Southern Ocean north of the sea ice (avg H11006 SD: 0.195 H11006 0.017 m H110021 ) (Figure 3), possibly due to lower rates of photobleaching or release of CDOM from the sea ice into the underlying seawater (Scully and Miller, 2000; Xie and Gosselin, 2005). However, we cannot ex- clude the possibility that the ship caused some release of CDOM from the ice during our passage through it. Absorption coeffi cients were also low within the Ross Sea during the development of the Phaeocystis antarctica bloom, even when surface waters were visibly green (chl a H11005 4? 8 ?g L H110021 ). For example, the absorption coeffi cient spectrum at station R14F was remarkably similar to that obtained in the Sargasso Sea during July 2004 (Figure 4), even though the chl a content of these two water samples differed by two orders of magnitude (7.0 versus 0.08 H9262g L H110021 , respectively). The main differences observed between the two spectra were seen at wavelengths less than approx- imately 350 nm, with the Sargasso Sea sample exhibiting a much higher spectral slope (e.g., S 275?295 of 0.0466 versus 0.0273 nm H110021 ) and the Ross Sea sample having a higher a ? between H11011300 and 350 nm. These differences are likely due to different autochthonous sources of CDOM as well as the presence of micromolar levels of nitrate in the Ant- arctic sample (and perhaps dissolved mycosporine amino acids (MAA) as well; see ?Ross Sea Temporal Trends in CDOM? section) compared to the low nanomolar levels of nitrate in the surface Sargasso Sea sample. FIGURE 3. Sea surface a 300 plotted as a function of degrees latitude south for the Oct?Nov 2005 transect from the Southern Ocean south of New Zealand to the Ross Sea. All data were obtained from water collected from the ship?s underway seawater pump system (intake at approximately 4-m depth) that was fi ltered inline through a 0.2-H9262m AS 75 Polycap fi lter. The dashed line represents the approximate location of the northern extent of seasonal sea ice in our transect. For details regarding the transect, see Kiene et al. (2007). FIGURE 4. Comparison of spectral absorption coeffi cients de- termined with seawater collected from 50 m in the Sargasso Sea (30?55.5H11032N, 65?17.8H11032W) on 21 July 2004 (dotted line) and from 40 m in the Ross Sea (77?31.2H11032S, 179?79.8H11032W) on 26 November 2005 (station R14B, solid line). Both samples were 0.2-H9262m grav- ity fi ltered prior to analysis. The chl a concentration was 0.08 and 7.0 H9262g L H110021 in the Sargasso Sea and Ross Sea, respectively. Absorp- tion spectra in the Sargasso Sea were determined with a Hewlett Packard (HP) 8453 UV-Vis photodiode array spectrophotometer equipped with a 5-cm path length rectangular microliter quartz fl ow cell; the Sargasso spectrum was referenced against Milli-Q water (for details, see Helms et al., 2008). The small discontinuity at 365 nm is an artifact associated with the HP spectrophotometer (Blough et al., 1993). 24_Kieber_pg319-334_Poles.indd 32424_Kieber_pg319-334_Poles.indd 324 11/18/08 9:15:31 AM11/18/08 9:15:31 AM CHROMOPHORIC DISSOLVED ORGANIC MATTER CYCLING 325 DOWNWELLING ATTENUATION COEFFICIENTS The optical clarity of the water column was reduced considerably from before the onset of the Phaeocystis ant- arctica bloom at station R10 to the end of the cruise at station R14. Before the onset of the Phaeocystis antarc- tica bloom (stations R10 and R13A? R13D), the photic zone exhibited a high degree of optical clarity, character- istic of type 1 open oceanic water (Mobley, 1994), with K d (?) decreasing exponentially in the UV with increasing wavelength and dominated by absorption by CDOM (Fig- ure 5). Prebloom K d (?) shown in Figure 5 for station R10 were nearly the same as observed at stations R13A? R13D (data not shown). For station R10, K d (?) ranged from 0.30 to 0.38 m H110021 at 305 nm, 0.18? 0.31 m H110021 at 320 nm, and 0.05? 0.15 m H110021 at 395 nm. Because of low biomass (chl a H11349 0.6 H9262g L H110021 ), the lowest values of K d (?), and therefore highest degree of optical clarity, were observed in the vis- ible between approximately 450 and 500 nm with K d (?) ranging from 0.05 to 0.11 m H110021 . When the bloom started to develop, the optical characteristics of the water, includ- ing the spectral structure of K d (?), changed dramatically. Over an approximately three-week period, values of K d (?) increased by a factor of six or more at some wavelengths, with peaks observed at two optical channels, 340 H11006 10 and 443 H11006 10 nm. At stations R13E and R14 (stations A through F) the bloom progressed to the point that the water column was more transparent to some wavelengths in the UV (e.g., 380 nm) relative to wavelengths in the blue portion of the solar spectrum (Figure 5). ROSS SEA TEMPORAL TRENDS IN CDOM Although UV and visible downwelling attenuation coeffi cients increased substantially when the Phaeocys- tis antarctica bloom was well developed in the Ross Sea (Figure 5), CDOM absorption coeffi cient spectra changed very little. To illustrate this point, an absorption coeffi cient spectrum of a 10-m sample taken before the onset of the bloom at station R10 was compared to a spectrum deter- mined for a 10-m sample at station R14 when the water was visibly green (Figure 6). As can be seen from com- parison of these two spectra, as well as from many other spectra not shown here, there was essentially no change in FIGURE 5. Spectral downwelling attenuation coeffi cients derived from PUV-2500 and PRR-600 profi les both before the Phaeocystis antarctica bloom (station R10D, circles) and during the development of the bloom (station R13E, triangles; station R14D, squares). Lines connecting the data do not represent a mathematical fi t of the data. Water column profi le cast locations are as follows: R10D, 13 No- vember 2005 New Zealand (NZ) time at 76?5.74H11032S, 170?15.70H11032W; R13E, 22 November 2005 NZ time at 77?5.20H11032S, 177?24.49H11032W; and R14D, 28 November 2005 NZ time at 77?3.61H11032S, 178?49.06H11032E. All optical profi les were conducted at approximately 1200 local noon NZ time. FIGURE 6. Spectral absorption coeffi cients plotted for 10-m samples from stations R10A (lower spectrum) and R14F (upper spectrum); latitude-longitude data for these stations are given in the Figure 8 caption. Thick lines represent the best fi t of the data from 275 to 295 nm and 350 to 400 nm, employing nonlinear regression analysis (equation 2). For station R10A, S 275?295 H11005 0.0280 H11006 0.0003 nm H110021 (r 2 H11005 0.997) and S 350?400 H11005 0.0115 H11006 0.0003 nm H110021 (r 2 H11005 0.904). For station R14F, S 275?295 H11005 0.0237 H11006 0.0004 nm H110021 (r 2 H11005 0.986) and S 350?400 H11005 0.0117 H11006 0.0005 nm H110021 (r 2 H11005 0.916). 24_Kieber_pg319-334_Poles.indd 32524_Kieber_pg319-334_Poles.indd 325 11/18/08 9:15:33 AM11/18/08 9:15:33 AM 326 SMITHSONIAN AT THE POLES / KIEBER, TOOLE, AND KIENE absorption coeffi cients or spectral slopes (S 350?400 ) at wave- lengths greater than approximately 350 nm. At shorter wavelengths, absorption coeffi cients and spectral slopes (S 275?295 ) varied by 15%? 30%, but with no consistent pat- tern among the many samples analyzed during a period when K d (340 and 443) and chl a changed by more than a factor of 6 and 100, respectively. Although there were generally only small changes in a ? spectra and S, there was an indication of a discernable increase in absorption coeffi cients in the vicinity of 330? 350 nm seen in a few sample spectra (station R14), with the presence of a small peak (or shoulder) noted in some cases. Previous results in the literature suggest that this peak may be due to the presence of MAA in the dissolved phase, possibly stemming from release by Phaeocystis ant- arctica, either through grazing, viral lysis, or direct release. The MAA are one of the primary UV-absorbing com- pounds detected in Phaeocystis antarctica (e.g., Riegger and Robinson, 1997; Moisan and Mitchell, 2001), and it would not be unreasonable for there to be some algal release of MAA into the dissolved phase. However, it is also possible that this UV absorption peak was an artifact of sample fi ltration (cf. Laurion et al., 2003). While arti- facts associated with sample fi ltration were not rigorously tested in this study, they are probably minimal because we prescreened water by gravity through 20-H9262m Nitex mesh to remove large aggregates and Phaeocystis colonies and then used gravity (hydrostatic) pressure for fi ltration through a 0.2-H9262m AS 75 POLYCAP fi lter. When gentle vacuum fi ltration was tested on samples collected during the bloom, we often observed a 330- to 350-nm peak that was not seen in CDOM spectra of the same samples that were prescreened and gravity fi ltered. Vacuum fi ltration is not recommended and may explain the presence of a peak in spectra for some samples that were analyzed during a bloom in Marguerite Bay along the Antarctic Peninsula (Patterson, 2000). The striking lack of change in CDOM spectra and spectral slopes during the development of the Phaeo- cystis antarctica bloom was also seen in temporal trends in surface a ? . Using 340 nm as an example, surface val- ues of a 340 in the upper 10 m did not appreciably change from 8 November (0.0782 m H110021 ) to 29 November (0.0837 m H110021 ), while over the same time frame, chl a changed by more than a factor of 100 from 0.084 to 8.45 H9262g L H110021 (Figure 7A). As with 340 nm, no signifi cant changes in a ? were observed at other wavelengths during the devel- opment of the bloom. Even though CDOM absorption coeffi cients did not change over time, downwelling at- tenuation coeffi cients increased dramatically (Figure 7B), FIGURE 7. Temporal trends in (A) surface chl a and a 340 and (B) K d (340) and a 340 from 8 November to 29 November 2005 (NZ time) in the Ross Sea, Antarctica. (C) Chlorophyll a plotted against K d (340). In Figure 7C, the line denotes the best fi t obtained from lin- ear correlation analysis: r 2 H11005 0.900, K d (340) H11005 0.053chl a H11001 0.128 paralleling increases in chl a (Figure 7C), especially in the vicinity of 340 and 443 nm (Figure 5), corresponding to particulate MAA and chl a absorption, respectively. The strong correlation between K d (340) and chl a shown in Figure 7C was also seen when K d (443) was plotted against chl a (r 2 H11005 0.937, data not shown). The nonzero 24_Kieber_pg319-334_Poles.indd 32624_Kieber_pg319-334_Poles.indd 326 11/18/08 9:15:34 AM11/18/08 9:15:34 AM CHROMOPHORIC DISSOLVED ORGANIC MATTER CYCLING 327 y-intercept in Figure 7C denotes the background K d (340) signal at very low chl a due to water, CDOM, and par- ticles in the Ross Sea. The lack of a clear temporal trend noted in a ? sur- face values during the Phaeocystis antarctica bloom in November 2005 was also evident in a ? and spectral slope depth profi les (Figures 8 and 9, respectively). Although we obtained multiple profi les at each station, results for only four CTD casts are shown in Figures 8 and 9 for clar- ity, with at least one CTD profi le depicted for each main hydrostation (R10, R13, and R14). The absorption coef- fi cient at 300 nm varied from approximately 0.18 to 0.30 m H110021 in the upper 100 m, with no temporal trend observed; some of the lowest and highest values were observed early and late in the cruise (Figure 8A). In the upper 50 m, later in the cruise, a 340 showed somewhat higher values at sta- tions R13E and R14F compared to stations R10 and R13A (Figure 8B), as did chl a (Figure 8C). However, while a 340 increased by 40%? 60%, chl a increased by more than two orders of magnitude. It was therefore not surprising that variations in a 340 did not correlate well with changes in chl a (r 2 H11005 0.384), as shown in Figure 8D. Spectral slopes also did not vary with any consistent trend. S 275-295 showed less than 25% variation over depth and time, ranging from H110110.023 to 0.031 nm H110021 (average 0.027 nm H110021 , n H11005 58, 7% RSD) during the cruise. Trends in S 350?400 were more variable (range 0.009? 0.017 nm H110021 , average 0.012 nm H110021 , n H11005 58, 15% RSD) but still showed no consistent trend with depth or time (Figure 9A and 9B). This variability was also seen in the ratio of the spec- tral slopes (Figure 9C), which showed no correlation to chl a (r 2 H11005 0.143; Figure 9D). FIGURE 8. Depth profi les of (A) absorption coeffi cient at 300 nm, (B) absorption coeffi cient at 340 nm, and (C) chl a. (D) Plot of a 340 versus chl a; linear correlation result is a 340 H11005 0.0046chl a H11001 0.091 with r 2 H11005 0.384. Symbols are: diamonds, station R10A (CTD cast at 0754 local NZ time on 10 November 2005; 76?13.85H11032S, 170?18.12H11032W); squares, station R13A (CTD cast at 0817 local NZ time on 18 November 2005; 77?35.16H11032S, 178?34.57H11032W); triangles, station R13E (CTD cast at 0823 local NZ time on 22 November 2005; 77?12.61H11032S, 177?23.73H11032W); and circles, station R14F (CTD cast at 0734 local NZ time on 30 November 2005; 77?15.35H11032S, 179?15.35H11032E). FIGURE 9. Depth profi les of (A) CDOM spectral slope from 275 to 295 nm, (B) CDOM spectral slope from 350 to 400 nm, and (C) the spectral slope ratio (S 275? 295 :S 350? 400 ). Horizontal error bars denote the standard deviation. (D) Spectral slope ratio (S R ) plotted against chl a; linear correlation results were S R H11005 H110020.0447chl a H11001 2.37, with r 2 H11005 0.143 (line not shown). For all four panels, symbols are: diamonds (station R10A), squares (station R13A), triangles (station R13E), and circles (station R14F). Cast times and station locations are given in Figure 8 caption. 24_Kieber_pg319-334_Poles.indd 32724_Kieber_pg319-334_Poles.indd 327 11/18/08 9:15:35 AM11/18/08 9:15:35 AM 328 SMITHSONIAN AT THE POLES / KIEBER, TOOLE, AND KIENE DISCUSSION CDOM absorption coeffi cients in the Southern Ocean and Ross Sea were consistently low throughout the cruise and along the transect south from New Zealand, with val- ues signifi cantly less than unity at wavelengths H11350300 nm (e.g., 0.15? 0.32 m H110021 at 300 nm). The a ? values determined in this study are slightly lower than values reported in the Weddell? Scotia Sea confl uence (Yocis et al., 2000) and along the Antarctic Peninsula (Sarpal et al., 1995; Yocis et al., 2000) (a 300 , range 0.19? 0.58 m H110021 ) and are similar to values obtained on a 2004? 2005 Ross Sea cruise (a 300 , range H110110.29? 0.32 m H110021 ) during the latter stages of the Phaeocystis antarctica bloom and transition to a diatom- dominated bloom (Kieber et al., 2007). These published fi ndings, along with results from our transect study and Ross Sea sampling, suggest that relatively low values of a ? are a general feature of Antarctic waters. By comparison, a 300 values in the Bering Strait? Chukchi Sea are higher, ranging from 0.3 to 2.1 m H110021 , with an average of 1.1 m H110021 (n H11005 62) (Ferenac, 2006). Similarly high a 300 values in the Greenland Sea (Stedmon and Markager, 2001) suggest that a ? values in the Antarctic are generally lower than those in Arctic waters, perhaps due to terrestrial inputs of DOM in the Arctic (Opsahl et al., 1999) that are lacking in the Antarctic. Although Ross Sea absorption coeffi cients were similar to those in the Sargasso Sea at longer wavelengths H11022350 nm, absorption coeffi cient spectra were different in the Ross Sea at shorter wavelengths (Figure 4), with higher a ? seen between approximately 290 and 350 nm. To illustrate this point further, we computed a 325 on all casts and depths (in the upper 140 m) during our cruise (n H11005 68) in order to directly compare them to results obtained by Nelson et al. (2007) in the Sargasso Sea. The a 325 values obtained during the Phaeocystis antarctica bloom were, on average (H11006 SD) 0.142 (H110060.017) and 0.153 (H110060.023) m H110021 (with an RSD of 12% and 15%) before and during the bloom, respectively, with no temporal trends noted. If we consider a 325 before the bloom (0.142 m H110021 ) and subtract the calculated contri- bution due to nitrate (present at 30 H9262M in the photic zone), then the average a 325 value due to CDOM is 0.120 m H110021 . This is approximately a factor of 2.5 higher than a 325 values reported by Nelson et al. (2007) in the surface Sargasso Sea (H110110.05 m H110021 at 325 nm) and reported by Morel et al. (2007) in the South Pacifi c gyre. While our a 325 values are higher than those Nelson et al. (2007) found in the Sargasso Sea, they are compa- rable to those Nelson et al. (2007) reported in the Subpo- lar Gyre (H110110.14? 0.26 m H110021 ), consistent with a slight pole- ward trend of increasing a 325 in open ocean waters due to a poleward decrease in CDOM photolysis rates and an increase in mixed layer depth (Nelson and Siegel, 2002). However, since not all wavelengths showed the same dif- ference between samples (see Figure 4; e.g., a ? H11021290 nm were smaller in the Ross Sea compared to the Sargasso Sea), care should be taken in extrapolating trends for all wavelengths given that there are likely regional differences in spectral shapes due to differences in CDOM source and removal pathways. The supposition that CDOM photobleaching rates are slow in Antarctic waters is supported by fi eld evi- dence. When 0.2-H9262m-fi ltered Ross Sea seawater samples in quartz tubes were exposed to sunlight for approximately eight hours, no measurable CDOM photobleaching was observed. This contrasts results from the Sargasso Sea or other tropical and temperate waters, where H1101110% loss in CDOM absorption coeffi cients is observed after six to eight hours of exposure to sunlight (D. J. Kieber, unpublished results). This difference is likely driven by lower actinic fl uxes at polar latitudes. However, differences in DOM reactivity cannot be ruled out. In addition to direct pho- tobleaching experiments, no evidence for photobleaching was observed in a CDOM depth profi les. At 300 nm, for ex- ample, absorption coeffi cients were uniform in the upper 100 m or showed a slight increase near the surface (Figure 8), a trend that is opposite of what would be expected if CDOM photolysis (bleaching) controlled near-surface a ? . It is also possible that mixing masked photobleaching, but this was not evaluated. ABSORPTION COEFFICIENT AT 443 nm Understanding what controls spatial and temporal trends in near-surface light attenuation is critical to accu- rately interpret remotely sensed ocean color data from the Sea-viewing Wide Field-of-view Sensor (SeaWiFS) or future missions (see Smith and Comiso, 2009, this volume). One of the primary limitations in using satellite ocean color data to model CDOM distributions is the lack of directly mea- sured CDOM spectra in the oceans for model calibration and validation, particularly in open oceanic environments and polar waters (Siegel et al., 2002). The 443 waveband corresponds to the chl a absorption peak and is often used in bio-optical algorithms for pigment concentrations, al- though model estimates suggest that CDOM and detrital absorption account globally for H1102250% of total nonwater absorption at this wavelength (Siegel et al., 2002). During 24_Kieber_pg319-334_Poles.indd 32824_Kieber_pg319-334_Poles.indd 328 11/18/08 9:15:36 AM11/18/08 9:15:36 AM CHROMOPHORIC DISSOLVED ORGANIC MATTER CYCLING 329 our cruise, chl a concentrations varied one-hundred-fold, while a 443 varied approximately 35% and showed no cor- relation to chl a. In the upper water column, a 443 was, on average (H11006 SD) 0.032 (H110060.011) m H110021 , with no differ- ence observed in prebloom values (0.031 H11006 0.011 m H110021 ) compared to values obtained during the bloom (0.033 H11006 0.012 m H110021 ). These observations fall within the range of wintertime zonally predicted values for southern latitudes between approximately 60? and 75?S (H110110.009? 0.05 m H110021 ) (Siegel et al., 2002) and suggest that the phytoplankton bloom is not directly responsible for the observed CDOM absorption. While we do not have direct observations to calculate the contribution of CDOM absorption to total absorp- tion, K d (?) values allow us to assess the contribution of CDOM to total light attenuation. K d (?) calculated from upper ocean optical profi les is the sum of component K d (?) from pure water, CDOM, and particulate material (phyto- plankton and detritus). The contribution from particulate material can be determined using K d (?) from pure water (Morel and Maritorena, 2001) in conjunction with mea- sured CDOM absorption coeffi cients. To a fi rst approxima- tion the majority of the particulate matter is expected to be living phytoplankton, as the Ross Sea is spatially removed from sources of terrigenous material. At the prebloom sta- tion R10A, the chl a concentration in the upper 30 m was, on average (H11006SD), 0.51 (H110060.04) H9262g L H110021 and K d (443) was relatively low (0.0684 m H110021 ). Pure water contributed 14.5% of the total attenuation at 443 nm while CDOM accounted for 42.8% (H110064.7%) (avg H11006 RSD) of the non water absorp- tion, with 57.2% (H110064.7%) accounted for by particles. At the bloom station R14D, the average chl a concentration was an order of magnitude greater (6.87 H11006 1.4 H9262g L H110021 ), and pure water contributed 2.1% of the total attenuation at 443 nm. Chromophoric dissolved organic matter accounted for only 3.5% (H110060.9%) of the total nonwater attenuation, with particles contributing the remaining 96.5% (H110060.9%). Not surprisingly, this confi rms that the attenuation of blue and green wavelengths was dominated by particles and not by dissolved constituents during the bloom. Many of the current suite of satellite algorithms (e.g., OC4v4, O?Reilly et al., 2000) have been shown to poorly predict chl a in the Southern Ocean, potentially because of the unique optical properties of large Phaeocystis antarc- tica colonies or the assumption that water column optical properties covary with chl a (e.g., Siegel et al., 2005). Our results confi rm that semianalytical approaches, which in- dividually solve for optical components, are necessary to describe the decoupling of CDOM and particulate mate- rial during Phaeocystis antarctica blooms in the Ross Sea and ultimately allow accurate chl a retrievals. SPECTRAL SLOPE The spectral slope (S) varies substantially in natural wa- ters, and it has been used to provide information about the source, structure, and history of CDOM (Blough and Del Vecchio, 2002). However, while spectral slopes have been reported in the literature for a range of natural waters, they are nearly impossible to compare because of differences in how S values have been determined and because different wavelength ranges have been considered (Twardowski et al., 2004). Indeed, when we employed a nonlinear fi tting routine to determine the spectral slope over several broad wavelength ranges, a range of S values was obtained for the same water sample [e.g., 0.0159 nm H110021 (290? 500 nm), 0.0124 nm H110021 (320? 500 nm), 0.0118 nm H110021 (340? 500 nm), 0.0133 nm H110021 (360? 500 nm), 0.0159 nm H110021 (380? 500 nm), 0.0164 nm H110021 (400? 500 nm)]. These variations in the slope indicate that Antaractic a ? spectra do not fi t a simple expo- nential function, as also observed by Sarpal et al. (1995) for samples along the Antarctic Peninsula. Therefore, instead of computing S over relatively broad wavelength ranges, we chose instead to compute S over two relatively narrow wavelength ranges proposed by Helms et al. (2008), 275? 295 nm and 350? 400 nm. These wavelength ranges were chosen because they can be determined with high precision and they are less prone to errors in how the data are mathe- matically fi t relative to broad wavelength ranges (e.g., 290? 700 nm; Helms et al., 2008). Additionally, the ratio, S R , for these two wavelength ranges should facilitate compari- son of different natural waters. In our study, the spectral slope between 275 and 295 nm was in all cases signifi cantly higher than at 350? 400 nm (H110110.028 nm H110021 versus H110110.013 nm H110021 , respectively), and S R was low, ranging between 1 and 3, with an average value of 2.32 (e.g., Figure 9C). Our S R are similar to coastal values observed in the Georgia Bight (avg H11006 SD: 1.75 H11006 0.15) and elsewhere even though a ? in coastal areas are much higher than we ob- served in the Ross Sea (e.g., H110221 m H110021 versus H110210.3 m H110021 at 300 nm, respectively). Likewise, the S R values we found in the Ross Sea are much lower than those observed in open oceanic sites, where values as high as 9 or more have been observed (Helms et al., 2008). Helms et al. (2008) found a strong positive correlation between S R and the relative proportion of low molecular weight DOM versus high mo- lecular weight (HMW) DOM in the sample. If the results of Helms et al. can be extrapolated to Antarctic waters, then 24_Kieber_pg319-334_Poles.indd 32924_Kieber_pg319-334_Poles.indd 329 11/18/08 9:15:36 AM11/18/08 9:15:36 AM 330 SMITHSONIAN AT THE POLES / KIEBER, TOOLE, AND KIENE our low S R results would suggest that the Ross Sea samples contained a relatively high proportion of HMW DOM compared to what is observed in oligotrophic waters. The relatively higher molecular weight DOM and low S R in the Ross Sea may be related to the lower photobleaching rates that are observed in the Antarctic, since photobleaching is the main mechanism to increase S R and remove HMW DOM (Helms et al., 2008). CDOM SOURCE The CDOM present in coastal areas near rivers and in estuaries in subtropical-temperate and northern boreal lat- itudes is predominantly of terrestrial origin (e.g., Blough et al., 1993; Blough and Del Vecchio, 2002; Rochelle-Newall and Fisher, 2002a), but in the open ocean, as in the Ant- arctic, terrigenous CDOM is only a minor component of the total DOM pool (Opsahl and Benner, 1997; Nelson and Siegel, 2002). Phytoplankton are expected to be the main source of CDOM in the open ocean, although they are not thought to be directly responsible for CDOM production (Bri- caud et al., 1981; Carder et al., 1989; Del Castillo et al., 2000; Nelson et al., 1998, 2004; Rochelle-Newall and Fisher 2002b; Ferenac, 2006). Our results are consistent with this fi nding (Figure 7). With no substantial grazing pressure (Rose and Caron, 2007, and as noted by our colleagues D. Caron and R. Gast during our cruise to the Ross Sea in early November 2005) and very low bacterial activity (Ducklow et al., 2001; Del Valle et al., in press), it is not surprising that CDOM a 300 and a 340 changed very little during the bloom. In fact, a 300 did not appar- ently increase even during the latter stages of the Ross Sea Phaeocystis antarctica bloom when the microbial ac- tivity was higher (Kieber et al., 2007: fi g. 5). This fi nd- ing is rather remarkable since DOC is known to increase by H1101150% during the Phaeocystis antarctica bloom from H1101142 to H1102260 H9262M in the Ross Sea (Carlson et al., 1998). We also observed a DOC increase during our cruise from background levels (H1101143 H9262M at 130 m) to 53-H9262M DOC near the surface (based on one DOC profi le obtained at our last station, R14F, on 30 November 2005). This dif- ference suggests that the DOC increase was due to the release of nonchromophoric material by Phaeocystis ant- arctica. Field results support this supposition, showing that the main DOC produced by Phaeocystis antarctica is carbohydrates (Mathot et al., 2000). The temporal and spatial decoupling between DOC and CDOM cycles in the Ross Sea photic zone indicate that CDOM was produced later in the season or elsewhere in the water column. Since evidence from Kieber et al. (2007) suggests that additional CDOM did not accumu- late later in the season, it was likely produced elsewhere. Previous studies have shown that sea ice is a rich source of CDOM (Scully and Miller, 2000; Xie and Gosselin, 2005). It is therefore possible that the pack ice along the edges of the Polynya was an important source of CDOM in the photic zone, especially the bottom of the ice, which was visibly light brown with ice algae. It is also possible that CDOM was produced below the photic zone and reached the photic layer via vertical mixing. Several lines of evi- dence indicate that Ross Sea Phaeocystis antarctica may be exported out of the photic zone (DiTullio et al., 2000; Rellinger et al., in press) and perhaps reach the ocean fl oor (H11011900 m) (DiTullio et al., 2000), as seen in the Arctic. Arctic algal blooms often occur in the early spring when the water is still too cold for signifi cant zooplankton graz- ing and bacterial growth (Overland and Stabeno, 2004). As a consequence, the algae sink out of the photic zone and settle to the ocean fl oor rather than being consumed, thereby providing a DOM source for long-term CDOM production in dark bottom waters. A similar phenomenon may occur in the Ross Sea. The possibility that CDOM may be generated in deep waters, with no photobleach- ing, might explain the high photosensitizing capacity of this CDOM for important reactions like DMS photolysis (Toole et al., 2004). Deepwater production of CDOM in the Ross Sea would explain why our a 325 values are comparable to those observed by Nelson et al. (2007: fi gs. 8? 9) in deep water (H110110.1? 0.2 m H110021 ) and nearly the same as would be predicted on the basis of values in Antarctic Bottom Water and Antarctic Intermediate Water (average of H110110.13? 0.14 m H110021 ). Although speculative, it is possible that CDOM is exported from the Southern Ocean to deep waters at temperate- subtropical latitudes, which would be consis- tent with CDOM as a tracer of oceanic circulation ( Nelson et al., 2007). CONCLUSIONS Despite the necessity of understanding the spatial and temporal distributions of CDOM for remote sens- ing, photochemical, and biogeochemical applications, few measurements have been made in the Southern Ocean. Our results indicate that Antarctic CDOM shows spectral properties that are intermediate between what is observed 24_Kieber_pg319-334_Poles.indd 33024_Kieber_pg319-334_Poles.indd 330 11/18/08 9:15:37 AM11/18/08 9:15:37 AM CHROMOPHORIC DISSOLVED ORGANIC MATTER CYCLING 331 in coastal environments and properties observed in the main oceanic gyres. We suggest this trend is largely due to slow photobleaching rates and shading from Phaeocystis antarctica and other bloom-forming species that contain substantial MAA. While CDOM spectral absorption coeffi cients are low in Antarctic waters, they are generally higher than surface water a ? in low-latitude, open-ocean waters, such as the Sargasso Sea, supporting the supposition of a pole- ward increase in a CDOM in the open ocean. Our results suggest that CDOM in the Ross Sea is not coupled di- rectly to algal production of organic matter in the photic zone. This indicates that case I bio-optical algorithms, in which all in-water constituents and the underwater light fi eld are modeled to covary with chl a (e.g., Morel and Maritorena, 2001), are inappropriate. The decou- pling of the phytoplankton bloom and CDOM dynam- ics indicates that CDOM is produced from sea ice or the microbial degradation of algal-derived dissolved organic matter that was exported out of the photic zone. Ross Sea CDOM absorption coeffi cients are similar in mag- nitude to values in Antarctic-infl uenced deep waters of the North Atlantic (Nelson et al., 2007), suggesting long- range transport of CDOM produced in the Ross Sea via Antarctic Intermediate and Bottom Water. ACKNOWLEDGMENTS This work was supported by the NSF (grant OPP- 0230499, DJK; grant OPP-0230497, RPK). Any opinions, fi ndings, and conclusions or recommendations expressed in this paper are those of the authors and do not necessar- ily refl ect the views of the NSF. The authors gratefully ac- knowledge the chief scientists for the October? December 2005 Ross Sea cruise, Wade Jeffery (University of West Florida) and Patrick Neale (Smithsonian Environmental Research Center). Thanks are also extended to Patrick Neale, Wade Jeffery, and their research groups for col- lection of the optics profi les, and the captain and crew of the Nathanial B. Palmer for technical assistance. We also thank Joaquim Goes (Bigelow Laboratory for Ocean Sciences), Helga do S. Gomes (Bigelow Laboratory for Ocean Sciences), Cristina Sobrino (Smithsonian Environ- mental Research Center), George Westby (State University of New York, College of Environmental Science and For- estry: SUNY-ESF), John Bisgrove (SUNY-ESF), Hyakubun Harada (Dauphin Island Sea Lab, University of South Alabama), Jennifer Meeks (Dauphin Island Sea Lab, Uni- versity of South Alabama), Jordan Brinkley (SUNY-ESF), and Daniela del Valle (Dauphin Island Sea Lab, University of South Alabama) for their technical help with sampling during this study. LITERATURE CITED Becquevort, S., and W. O. Smith Jr. 2001. Aggregation, Sedimentation and Biodegradability of Phytoplankton-Derived Material During Spring in the Ross Sea, Antarctica. Deep-Sea Research, Part II, 48: 4155? 4178. Blough, N. V., and R. Del Vecchio. 2002. ?Chromophoric DOM in the Coastal Environment.? In Biogeochemistry of Marine Dissolved Organic Matter, ed. D. A. Hansell and C. A. Carlson, pp. 509? 546. San Diego: Academic Press. 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Kieber, and K. Mopper. 2000. Photochemical Produc- tion of Hydrogen Peroxide in Antarctic Waters. Deep-Sea Research, Part I 47: 1077? 1099. 24_Kieber_pg319-334_Poles.indd 33324_Kieber_pg319-334_Poles.indd 333 11/18/08 9:15:39 AM11/18/08 9:15:39 AM 24_Kieber_pg319-334_Poles.indd 33424_Kieber_pg319-334_Poles.indd 334 11/18/08 9:15:39 AM11/18/08 9:15:39 AM ABSTRACT. Weddell seals, like many true seals (Phocidae), store nutrients in body tis- sues prior to lactation and then expend these ?capital reserves? in pup rearing. During lactation, 40% or more of the initial mass of a lactating Weddell seal may be expended to cover the combined costs of maternal metabolism and milk production. However, most lactating Weddell seals also begin active diving to depths of 300 m or more by three to four weeks postpartum, and dietary biomarker data indicate that at least 70% of Weddell seals forage in late lactation. Thus, Weddell seals may employ a combined capital and income (foraging) strategy. Determining the relative importance of capital expenditures and food consumption to maternal reproduction will require accurate measurement of maternal energy expenditure, the magnitude of milk production, changes of maternal nutrient stores over lactation and the success of foraging efforts. Alternative scenarios include the following: (1) prey consumption is opportunistic rather than essential because body reserves of Weddell seals are suffi cient for reproduction, (2) foraging is necessary only in those females (such as small or young seals) that have limited body stores relative to lactation costs, and (3) successful foraging is critical to the lactation strategy of this species. If alternative 2 or 3 is correct, the drops in pup production observed in Erebus Bay (McMurdo Sound, Ross Sea) during years of unusually heavy ice accumulation may refl ect changes in foraging opportunities due to adverse impacts of heavy ice on primary production and on prey populations. Further study is needed on the effects of annual, cyclic, or long-term changes in prey abundance on Weddell seal reproduction. INTRODUCTION Mammalian reproduction is characterized by a period of lactation in which large quantities of nutrients are transferred from mother to young (Oftedal, 1984b). This process puts a great physiologic demand on the mother, who must either acquire the additional nutrients needed for milk secretion by increased food consumption, mobilize nutrients from stored reserves in the body, or em- ploy some combination of both (Oftedal, 2000). Along the continuum from in- tensive foraging to sole dependence on stored reserves, mammals that rely mostly on feeding can be characterized as ?income breeders,? whereas those that rely on stored reserves are ?capital breeders? (J?nsson, 1997). Income breeders are highly infl uenced by local climatic conditions that impact immediate food sup- ply, whereas capital breeders should be relatively independent of food resources Regina Eisert Conservation Ecology Center, National Zoological Park, Smithsonian Institu- tion, P.O. Box 37012, MRC 5507, Washington, DC 20013-7012, USA. Olav T. Oftedal, Smith- sonian Environmental Research Center, 647 Con- tees Wharf Road, Edgewater, MD 21037, USA. Corresponding author: R. Eisert (eisertr@si.edu). Accepted 28 May 2008. Capital Expenditure and Income (Foraging) during Pinniped Lactation: The Example of the Weddell Seal (Leptonychotes weddellii) Regina Eisert and Olav T. Oftedal 25_Eisert_pg335-346_Poles.indd 33525_Eisert_pg335-346_Poles.indd 335 11/17/08 9:18:07 AM11/17/08 9:18:07 AM 336 SMITHSONIAN AT THE POLES / EISERT AND OFTEDAL during lactation by virtue of their previously stored body reserves. Among Antarctic mammals, two groups rely heavily on stored reserves during lactation: baleen whales (sub- order Mysticeti) and true seals (family Phocidae). Baleen whales, such as the blue whale (Balaenoptera musculus), fi n whale (Balaenoptera physalus), humpback whale (Megap- tera novaeangliae), and minke whale (Balaenoptera acu- torostrata), migrate to Antarctic waters to forage on sea- sonal abundances of prey, such as krill, squid, and fi sh, and deposit large amounts of fat and other body constituents at this time (Lockyer, 1981, 1984; Oftedal, 1997). How- ever, these baleen whales migrate back to subtropical or temperate regions to give birth and lactate. Stored energy and nutrients fuel most or all of lactation as these species feed little if at all at the calving grounds (Oftedal, 1997). Thus, baleen whales export substantial quantities of nutri- ents from the Southern Ocean to more temperate regions. By contrast, phocid seals, such as southern elephant seal (Mirounga leonina), crabeater seal (Lobodon carcinopha- gus), Ross seal (Ommatophoca rossii), leopard seal (Hy- drurga leptonyx), and Weddell seal (Leptonychotes wed- dellii), both feed and lactate in Antarctic areas. Elephant seals typically remain on land and fast throughout a three to four week lactation (Arnbom et al., 1997) and are thus true capital breeders. On the basis of data from satellite- linked dive recorders, Ross seals are also capital breeders, as they haul out on pack ice for only about 13 days in mid-November to give birth and lactate (Blix and Nord?y, 2007). Unfortunately, little is known about reproduction in crabeater or leopard seals, but Weddell seals appear to employ a hybrid breeding approach: partly capital use and partly food consumption. The ability to rely solely on stored reserves to support the energy and substrate demands of lactation is limited by body size (Oftedal, 2000). Nutrient reserves increase in di- rect proportion to body mass (BM 1.0 ), but rates of energy expenditure (including lactation) increase in proportion to body mass raised to the power of 3/4 (BM 0.75 ). Thus, the capacity to support metabolism and lactation from body stores alone increases with body size, and larger species can support metabolism and lactation from stored reserves for longer periods of time. The benefi t of being able to store large quantities of nutrients for subsequent use was likely an important factor in the evolution of large body size in both seals and whales. At 400? 500 kg, the female Weddell seal is one of the largest of the phocid seals and has long been assumed to rely on stored reserves for lactation (Tedman and Green, 1987). If so, lactating Weddell seals should be relatively immune to environmental variables that affect local food supply in the areas where they give birth and lactate. However, population censuses have indicated tremendous variation (H1102250%) in annual pup production associated with changes in ice conditions in Erebus Bay in the Ross Sea (R. Garrott, Montana State University, personal com- munication, 2007). It is not known whether this variation is related to ice-related changes in prey abundance and diversity or to some other consequence of sea ice accumu- lation, such as navigational diffi culties for seals traveling under the ice. A step in addressing this issue is to evalu- ate the importance of stored reserves versus acquisition of food to lactation performance of the Weddell seal. In this paper we briefl y discuss breeding strategy, mass change, lactation performance, and foraging by Weddell seals, with comparisons to other phocid species. This pa- per is a preliminary contribution based on a project in 2006? 2007 examining energy expenditure, milk produc- tion, and changes in body reserves in lactating Weddell seals in McMurdo Sound, Antarctica. EVOLUTION OF CAPITAL BREEDING AMONG PHOCID SEALS Animals that employ capital breeding incur energetic costs associated with the deposition, transport, and mobi- lization of stores (J?nsson, 1997). The resulting energetic ineffi ciency is thought to favor ?income breeding? except in specifi c circumstances such as uncertainty or inade- quacy of food at the time of reproduction (J?nsson, 1997). However, among some capital breeders, such as phocid seals that fast during lactation, a major benefi t appears to be abbreviation of lactation, with consequent reduc- tion of maternal metabolic overhead and the time devoted to pup rearing. Milk production from stored reserves is also much more effi cient than production based on food consumption (Agricultural Research Council, 1980), es- pecially if foraging requires signifi cant effort. This permits an increase in the proportion of energy available for trans- fer to the offspring (Fedak and Anderson, 1982; Costa et al., 1986). At the extreme, lactation is reduced to as little as four days in the hooded seal, with up to 88% of the energy transferred to pups incorporated into tissue growth (Bowen et al., 1985; Oftedal et al., 1993). Thus, it is not clear that capital breeding is always more energetically costly than income breeding. A variety of other param- eters, including animal size, food availability, transport costs, neonatal developmental state, and type of maternal care, are thought to be important to the evolution of capi- 25_Eisert_pg335-346_Poles.indd 33625_Eisert_pg335-346_Poles.indd 336 11/17/08 9:18:07 AM11/17/08 9:18:07 AM WEDDELL SEAL CAPITAL BALANCE DURING LACTATION 337 tal breeding systems (Boyd, 1998; Trillmich and Weissing, 2006; Houston et al., 2006). There is also uncertainty whether maternal capital expenditure is limited primarily by energy or by nutrient stores, such as protein. In the fasting state, catabolized pro- tein is lost continually from the body (Nord?y et al., 1990; Owen et al., 1998), and lactating mammals must export milk protein to support offspring growth. Yet excessive loss of body protein leads to progressive and eventually lethal loss of function (Oftedal, 1993; Liu and Barrett, 2002). Animals that fast during lactation typically pro- duce milks that are low in both protein and carbohydrate (Oftedal, 1993). As both protein and carbohydrate in milk potentially derive from amino acids (either directly or via gluconeogenesis), this suggests that high protein demands may be selected against during the evolution of capital breeding (Oftedal, 1993). In the grey seal (Halichoerus grypus), daily milk production and fi nal offspring mass were signifi cantly correlated with initial maternal protein but not initial fat stores (Mellish et al., 1999a), despite the fact that most of maternal body energy reserves are stored as fat. Although phocid seals are often thought to be un- usually effi cient at conserving protein during fasting, this assumption may have to be reconsidered (Eisert, 2003). Thus, capital breeding may be limited by the size of pro- tein stores as well as by the magnitude of energy stores. Seals are the best-studied group of mammalian capital breeders (Oftedal et al., 1987a; Costa, 1991; Boness and Bowen, 1996; Boyd, 1998; Mellish et al., 2000; Oftedal, 2000; Schulz and Bowen, 2004). Otariid seals remain ashore for approximately one week after giving birth and transfer approximately 4% of body protein and 12% of body energy to their pups, after which they undertake regu- lar foraging trips to sea (Oftedal et al., 1987a; Costa, 1991; Oftedal, 2000). This strategy of an initial fasting period followed by foraging cycles occurs in at least one phocid, the harbor seal Phoca vitulina (Boness et al., 1994), and perhaps in other species that feed during lactation [e.g., bearded seal Erignathus barbatus, harp seal P. groenland- ica, and ringed seal P. hispida (Lydersen and Kovacs, 1996, 1999)]. However, many large phocids fast throughout the lactation period [e.g., land-breeding grey seal H. grypus, hooded seal Cystophora cristata and elephant seals Mir- ounga angustirostris and M. leonina (Fedak and Anderson, 1982; Costa et al., 1986; Oftedal et al., 1993; Arnbom et al., 1997)]. As the true seals (family Phocidae) encompass a wide spectrum from mixed capital-income to extreme capi- tal breeding, this family is an excellent model system for testing hypotheses about the evolution of capital breeding strategies. Factors thought to have favored the evolution of extreme capital breeding in phocids include large body size (Boness and Bowen, 1996; Oftedal, 2000), limited avail- ability of food (Boyd, 1998), the impact of unstable nurs- ing substrates (Oftedal et al., 1987a; Lydersen and Kovacs, 1999), and reduction of maternal metabolic overhead costs (Fedak and Anderson, 1982; Costa, 1991). WEDDELL SEAL: EXAMPLE OF AN INTERMEDIATE STRATEGY? The Weddell seal represents an interesting, if not fully understood, example of a species where a continuum of capital to mixed capital-income breeding strategies may occur within the same population. Lactating Weddell fe- males fast and remain with their offspring for at least the fi rst week postpartum, but on the basis of a new biomarker method of detecting feeding (Eisert et al., 2005), at least 70% of females feed to some extent during the latter half of a lactation period that lasts six to eight weeks (Bertram, 1940; Kaufmann et al., 1975; Thomas and DeMaster, 1983). During late lactation, an increase in diving activity (Hindell et al., 1999; Sato et al., 2002) and a decrease in rates of maternal mass loss relative to pup mass gain have also been observed (Hill, 1987; Testa et al., 1989). How- ever, the importance of food intake to the energy and nu- trient budgets or to reproductive success of lactating Wed- dell seals is not known, nor has the magnitude of capital expenditure (depletion of maternal body stores) been stud- ied. Three scenarios appear possible: (1) Females are able to complete lactation without food intake but take prey opportunistically (until recently, the prevailing belief). (2) Because of individual differences in nutrient stores and re- productive demand, some females (such as small or young females) have an obligatory need for food intake, while others do not. (3) Food intake is an essential part of the lactation strategy of this species because maternal body stores are inadequate in the face of such an extended lacta- tion period (the longest of any phocid). Uncertainty regarding the dependency of lactating Weddell seals on local food resources complicates efforts to interpret the infl uence of environmental factors on ma- ternal condition (Hill, 1987; Hastings and Testa, 1998), pup growth and survival (Bryden et al., 1984; Tedman, 1985; Tedman and Green, 1987; Testa et al., 1989; Burns and Testa, 1997), and population dynamics (Stirling, 1967; Siniff et al., 1977; Testa, 1987; Hastings and Testa, 1998). A strong dependency, in some or all females, on local food resources for successful lactation might limit breed- ing colonies to areas of local prey abundance or result in 25_Eisert_pg335-346_Poles.indd 33725_Eisert_pg335-346_Poles.indd 337 11/17/08 9:18:08 AM11/17/08 9:18:08 AM 338 SMITHSONIAN AT THE POLES / EISERT AND OFTEDAL the vulnerability of populations to annual or long-term changes in prey availability, as might occur due to changes in sea ice or shifts in water currents. MASS CHANGES DURING WEDDELL SEAL LACTATION Prior work on Weddell seals has focused primarily on mass changes of mothers and their pups, under the as- sumption that if mothers are fasting, there should be cor- respondence between maternal mass loss, maternal milk output, and pup mass gain, as is the case in other true seals that fast throughout lactation. In these species, maternal body mass and age are strong determinants of total milk energy output and, consequently, of pup growth and wean- ing mass (Iverson et al., 1993; Fedak et al., 1996; Arnbom et al., 1997; Mellish et al., 1999b). By contrast, females feed during a variable proportion of lactation in almost half of extant phocid species (Bonner, 1984; Oftedal et al., 1987a; Boness et al., 1994; Boness and Bowen, 1996; Lydersen and Kovacs, 1999; Eisert, 2003). Bowen et al. (2001a) found that the positive correlation of maternal body mass with pup weaning mass was much weaker in harbor seals than in species that fast during lactation, pre- sumably because supplementary feeding results in a par- tial decoupling of maternal mass loss and milk transfer to the pup. Similar patterns have been found in ice-breeding grey seals H. grypus and harp seals P. groenlandica (Baker et al., 1995; Lydersen and Kovacs, 1996, 1999). Extant data for Weddell seals are more complex. Wed- dell seal females certainly lose a large amount of body mass: for example, females that we studied in 2006 and 2007 lost 40% of their two-day postpartum mass during about 40 days lactation (Figure 1). The daily mass loss of 1.0% of initial mass is lower than values of 1.5%? 3.4% for fasting and lactating females of the northern elephant seal, south- ern elephant seal, land-breeding gray seal, and hooded seal (Costa et al., 1986; Carlini et al., 1997; Mellish et al., 1999a, 1999b), but Weddell seal lactation is so prolonged that overall mass loss (42%) is equal to or greater than that in the other species (14%? 39%). If mass loss is standard- ized to a lactation length of 42 days, initial mass predicts 66% of the variation in mass loss, indicating that large fe- males lose more mass than small females (Figure 2). Is this because large females expend more energy (on metabolism and milk production) or because they feed less? Females that lose more mass also support more mass gain by their pups: pup mass gain was positively correlated to maternal mass loss (Figure 3). Tedman and Green (1987) found a similar strong positive correlation (r H11005 0.85, P H11021 0.001) between maternal mass loss and pup mass gain, whereas data from studies by Hill (1987) and Testa et al. (1989) indicate a much weaker correlation between maternal mass loss and pup mass gain (r H11005 0.16, P H11005 0.005, n H11005 35). FIGURE 1. Change in body mass of lactating Weddell seals from early (2? 3 days postpartum) to late (35? 45 days postpartum) lac- tation at Hutton Cliffs, Erebus Bay, McMurdo Sound. Data were obtained from 24 females in 2006 and 2007. Average rate of mass loss was 1.0% of initial mass per day. FIGURE 2. Relationship of maternal mass loss to initial maternal mass of lactating Weddell seals. Initial mass was measured at two to three days postpartum. Mass loss was normalized to 42 days and compared to initial mass by Deming linear regression. Data are from the same 24 females at Hutton Cliffs, Erebus Bay, McMurdo Sound, as in Figure 1. 25_Eisert_pg335-346_Poles.indd 33825_Eisert_pg335-346_Poles.indd 338 11/17/08 9:18:08 AM11/17/08 9:18:08 AM WEDDELL SEAL CAPITAL BALANCE DURING LACTATION 339 This difference could stem from differences in the masses of animals studied: in our study and in that of Tedman and Green (1987) mean female mass was about 450 kg at the beginning of lactation, whereas the average in Hill?s study was 406 kg. This suggests that the strength of the correlation of maternal mass loss and pup mass gain may increase with maternal size. Assuming that decoupling of maternal mass loss and pup growth in the Weddell seal can be attributed to foraging, feeding may be obligatory for small females but optional or opportunistic for large females (Testa et al., 1989). ISOTOPIC MEASUREMENTS OF EXPENDITURES Change in body mass alone is, at best, an imprecise measure of energy expenditure (Blaxter, 1989) and is in- valid if animals are obtaining signifi cant energy from food. The very high costs of lactation entail both metabolic costs (such as the energy expenditure associated with maternal attendance of pups and the energetic cost of milk synthesis) and substrate costs (the energy transferred into milk as fat, protein, carbohydrate, and minor constituents). Currently, the only method of accurately assessing metabolic energy ex- penditure in wild animals is the doubly labeled water (DLW) technique in which differences in the kinetics of hydrogen and oxygen isotopes provide an estimate of carbon diox- ide production (see reviews by Nagy, 1980, and Speakman, 1997). Because of the high economic cost of 18 O-labeled water, this procedure has usually been applied to mammals of small body size (Speakman, 1997); among phocids it has been applied to pups (e.g., Kretzmann et al., 1993; Lydersen and Kovacs, 1996). However, the DLW method may pro- vide valuable insight into maternal metabolic expenditures during lactation, as reported for sea lions and fur seals (e.g., Arnould et al., 1996; Costa and Gales, 2000, 2003). In Weddell seals it would be particularly interesting to know if metabolic energy expenditures vary in accord with diving activity, stage of lactation, and food consumption. In ani- mals that fast, one would expect metabolic rates to decline over the course of the fast, whereas energy expenditures should increase with increases in activity and in association with digestion and metabolism of food constituents (e.g., Blaxter, 1989; Speakman, 1997). However, there remain a number of technical issues to overcome, including selection of an appropriate model of isotope behavior and estima- tion of average respiratory quotient (RQ, ratio of carbon dioxide production to oxygen consumption), which differs between fasting and feeding animals. Model and RQ errors can directly impact energetic estimates and thus need to be assessed ( Speakman, 1997). There are also logistic prob- lems in accurately administering water isotopes to large, unsedated animals living in very cold and windy environ- ments, but these are not prohibitive: in 2006 and 2007, we successfully dosed about 20 lactating Weddell seals in Erebus Bay, McMurdo Sound, with doubly labeled water; sample analyses are still in progress. The DLW method does not, however, measure export of substrates via milk. It is therefore also necessary to measure milk yield and milk composition to estimate re- productive costs associated with the output of milk con- stituents (Oftedal, 1984b). The most widely used method for estimating milk production in seals relies on the dilu- tion of hydrogen isotope? labeled water in nursing young (e.g., Costa et al., 1986; Oftedal and Iverson, 1987; Oftedal et al., 1987b; Tedman and Green, 1987; Lydersen et al., 1992; Iverson et al., 1993; Lydersen and Ham- mill, 1993; Oftedal et al., 1993; Lydersen and Kovacs, 1996; Oftedal et al., 1996; Lydersen et al., 1997; Mellish et al., 1999a; Arnould and Hindell, 2002). If milk is the exclusive source of water (both free and metabolic) for the offspring, then milk consumption can be estimated from water turnover and milk composition (Oftedal and Iverson, 1987). The accuracy of this method depends on the ability to correct estimates of milk intake for isotope recycling (Baverstock and Green, 1975; Oftedal, 1984a), for changes in pool size (Dove and Freer, 1979; Oftedal, FIGURE 3. Pup mass gain in relation to maternal mass loss of lac- tating Weddell seals. Mother and pup data are paired (n H11005 24) and refl ect the same time periods (from 2? 3 days to 35? 45 days post- partum) for both. Pup gain and maternal loss were compared by Deming linear regression. 25_Eisert_pg335-346_Poles.indd 33925_Eisert_pg335-346_Poles.indd 339 11/17/08 9:18:16 AM11/17/08 9:18:16 AM 340 SMITHSONIAN AT THE POLES / EISERT AND OFTEDAL 1984a), and for any water obtained by offspring from sources other than milk (Holleman et al., 1975; Dove, 1988), such as consumption of prey, snow, or seawater and metabolic water production. Isotope studies have demonstrated that milk energy output in seals is inversely proportional to lactation length: seals with very short lactations, such as species that breed on unstable pack ice, have much higher daily energy outputs than species that breed on stable substrates, such as land and fast ice (Oftedal et al., 1987a). The sole published attempt at measuring milk pro- duction in Weddell seals employed two isotopes ( 2 H and 22 Na) to determine if pups were ingesting water from sources other than milk (Tedman and Green, 1987). Tedman and Green argued that if pups were obtaining all or most of their water from milk, the sodium intake predicted from milk consumption (calculated milk in- take from 2 H turnover multiplied by milk sodium con- tent) would be similar to that estimated from turnover of 22 Na. As the observed discrepancy was not large, they concluded that intake of seawater or sodium-containing prey must have been minor (Tedman and Green, 1987). However, large (20%) underestimation of sodium intake occurs in 22 Na turnover measurements on suckling young (Green and Newgrain, 1979), and Tedman and Green (1987) do not state whether this error was corrected for in their study. The potential importance of nonmilk water as a confounding effect in isotope studies warrants further study, especially as Weddell pups have been ob- served to grab snow in their mouths and may consume it. At present, the Tedman and Green (1987) data are the only published data on Weddell seal lactation, and the estimated milk yield of about 3.5 kg/d or 160 kg milk over the lactation period has been cited repeatedly in comparative studies (e.g., Oftedal et al., 1987a, 1996; Costa, 1991; Boness and Bowen, 1996; Oftedal, 2000) but is in need of reevaluation. In order to avoid possible errors caused by consump- tion of food or water by nursing Weddell seal pups, we recommend use of a two-hydrogen isotope method origi- nally developed for terrestrial herbivores in which the suckling young begin to feed on solid foods during lacta- tion (e.g., Holleman et al., 1975, 1988; Wright and Wolff, 1976; Oftedal, 1981; Dove, 1988; Carl and Robbins, 1988; Reese and Robbins, 1994). Water containing tri- tium ( 3 HHO) is given to the mother and water containing deuterium ( 2 H 2 O) is given to offspring so that their body water pools are separately labeled. Thus, water turnover in mother and offspring can be measured independently. In the pup, tritium concentrations rise so long as tritium intake (via milk water) exceeds tritium loss (via excre- tion), reaching a plateau when intake and loss are equal. In a Weddell seal pup this occurs after about two weeks (Figure 4). As tritium loss can be estimated from the rate of water turnover of the pup (measured from deuterium kinetics) and tritium intake equals milk tritium concentra- tion multiplied by milk water intake, modeling of isotope fl uxes allows calculation of mean water intake from milk. Unlike single isotope methods, this procedure allows milk water intake by the pup to be distinguished from infl ux of water from all other sources, such as drinking, feeding, and metabolic water production. Once milk water intake is known, milk production can be calculated from milk composition. We applied this dual-isotope method on about 20 Weddell pups in Erebus Bay, McMurdo Sound, in 2006 and 2007. FIGURE 4. Illustration of the dual-isotope procedure for a Weddell seal mother and her pup. The mother was administered tritium- labeled ( 3 HHO) water and the pup was given deuterium-labeled water ( 2 H 2 0) by intravenous infusion at the times indicated by verti- cal arrows. While deuterium levels ( 2 H 2 0) declined following admin- istration, tritium levels ( 3 HHO) in the pup rose towards a plateau level, indicating that tritium intake via milk water exceeded tritium losses of the pup over this time period. Deuterium was measured by isotope ratio mass spectrometry, and tritium was measured by scintillation counting. Mathematical modeling of isotope behavior indicated that milk intake was about 2.5 kg over this period. 25_Eisert_pg335-346_Poles.indd 34025_Eisert_pg335-346_Poles.indd 340 11/17/08 9:18:17 AM11/17/08 9:18:17 AM WEDDELL SEAL CAPITAL BALANCE DURING LACTATION 341 MONITORING FOOD CONSUMPTION DURING THE LACTATION PERIOD If Weddell seals did not enter the water during lacta- tion, it would be obvious that they must be fasting, but this is not the case. Weddell seal mothers usually remain on the ice with their pups for the fi rst one to two weeks of lactation, but then most mothers begin diving bouts that typically increase in frequency and length as lactation proceeds. The diving behavior of Weddell seals, including lactating females as well as nursing, weaned, and yearling animals, has been extensively investigated with time-depth recorders, or TDRs, including satellite-uplinked instru- ments (e.g., Kooyman, 1967; Testa et al., 1989; Burns et al., 1997, 1999; Castellini et al., 1992; Sato et al., 2002, 2003; Williams et al., 2004; Fuiman et al., 2007). The recent development of instrumentation that records directional information ( Harcourt et al., 2000; Davis et al., 2001; Hindell et al., 2002; Mitani et al., 2003) allows examina- tion of dive behavior in three-dimensional space. Although food intake requires diving, even deep diving need not en- tail food consumption. In other words, one cannot equate dive records to actual food intake unless food intake can be independently confi rmed (Testa et al., 1989). This has led to development of a number of different methods to monitor food intake. Classically, the diet of the Weddell seal has been examined by stomach content analysis (Bertram, 1940; Dearborn, 1965; Pl?tz, 1986; Pl?tz et al., 1991), but lethal methods are no longer em- ployed, and gastric lavage of adults requires extensive restraint or chemical immobilization. Scat analysis can provide information on those prey that have identifi able indigestible parts (Burns et al., 1998; Lake et al., 2003) or even residual prey DNA (Casper et al., 2007) in the scats. However, the relative proportions of prey are diffi - cult to quantify without extensive feeding trials to develop correction factors for the differential rates of digestion of prey. This is not feasible in free-ranging Weddell seals. In addition, the identity of the animal producing the scat and the time it was produced are often unknown. Occasion- ally, Weddell seals are observed to bring large prey, such as Antarctic toothfi sh (Dissostichus mawsoni) to holes in the ice (e.g., Caelhaem and Christoffel, 1969). Such ob- servations can be extended by deploying animal-borne underwater cameras (Marshall, 1998; Davis et al., 1999; Bowen et al., 2002; Fuiman et al., 2002; Sato et al., 2002, 2003; Fuiman et al., 2007). Although dives in which po- tential prey are visible have been termed ?foraging dives? ( Fuiman et al. 2007), it is diffi cult to determine the success of prey capture attempts from the images obtained. Of course, the images provide valuable information on hunt- ing methods of seals at depth. Another approach is to attach instruments in the mouth or digestive tract of seals to record feeding events. Sensors have been glued to the jaw of the seal that detect opening of the mouth (Bornemann et al., 1992; Pl?tz et al., 2001), but prey capture may be diffi cult to distinguish from other jaw movements during social behavior (e.g., threats and bites). Temperature-transmitting thermistors have been introduced into the stomach to monitor changes in the temperature of stomach contents associated with in- gestion of cold items, such as ectothermic fi sh ( Bornemann, 1994; Hedd et al., 1996; Austin et al., 2006a, 2006b; Kuhn and Costa, 2006). However, stomach temperature is also affected by other factors such as water ingestion, gastric blood fl ow, thermal mass of gastric contents and therm- istor location, so that validation studies are essential to interpretation (Ponganis, 2007). Instruments may also re- quire considerable intervention to attach (e.g., prolonged anesthesia), may alter animal behavior, or may require re- captures for data acquisition. Thus, there is still a need for a simple method of determining when food is actually consumed by free-living seals. As an alternative approach, food energy intake of seals has been estimated from changes in whole-body water fl ux using isotope-labeled water (Costa, 1987; Bowen et al., 2001b). This method relies on the assumption that uptake of water from sources other than food (e.g., drinking) is minimal (Costa, 1987; Bowen et al., 2001b), yet seals are known to voluntarily consume both freshwater and seawater (Skalstad and Nord?y, 2000; Lea et al., 2002). Lactating Weddell seals have been observed to eat snow (Eisert and Oftedal, unpublished observations). This may lead to errors of unknown magnitude in both the detection and quantitation of food intake. As a result of the diffi culties in detecting and quantify- ing food intake, the energetics of lactation in the Weddell seal and in other species that feed during lactation are not well described (Schulz and Bowen, 2004). However, this may be improved by combining isotope methodology with new techniques for detecting feeding using biomarkers (Eisert et al., 2005). This approach allows food intake to be confi rmed at a specifi c point in time from the presence in body fl uids of dietary biomarkers, i.e., specifi c compounds that are absorbed intact from prey but are not generated by normal metabolic processes in the predator. On the ba- sis of studies in Weddell seals, we have identifi ed two suit- able compounds, arsenobetaine (AsB) and trimethylamine 25_Eisert_pg335-346_Poles.indd 34125_Eisert_pg335-346_Poles.indd 341 11/17/08 9:18:20 AM11/17/08 9:18:20 AM 342 SMITHSONIAN AT THE POLES / EISERT AND OFTEDAL N-oxide (TMAO) (Eisert, 2003; Eisert et al., 2005). Both are specifi c to, and apparently ubiquitous in, marine prey yet are neither stored nor synthesized by higher vertebrates and in mammals are eliminated rapidly from the circula- tion following ingestion (Edmonds and Francesconi, 1977, 1987, 1988; Yancey et al., 1982; Vahter et al., 1983; Al-Waiz et al., 1987, 1992; Van Waarde, 1988; Cullen and Reimer, 1989; Brown et al., 1990; Shibata et al., 1992; Smith et al., 1994; Svensson et al., 1994; Zhang et al., 1999; Lehmann et al., 2001). The biomarker method provides information on recent food intake within a timescale of hours to days, in contrast to fatty acid signatures or stable isotopes in fl uids or tissue samples, which integrate food intake over a period of months (Iverson et al., 1997a, 1997b; Brown et al., 1999). Investigations of the incidence of foraging in lactating Weddell seals using the biomarker method (Eisert et al., 2005) revealed that (1) ~70% of females studied in late lactation (H1102227 days postpartum) had concentrations of AsB and TMAO indicative of recent food intake, (2) most females appear to fast for the fi rst three to four weeks of lactation, in agreement with observed dive activity ( Hindell et al., 2002), and (3) feeding may commence as early as eight to nine days postpartum in some females. These re- sults suggest that the onset of feeding may vary substan- tially among lactating seals, and the possibility remains that some females fast throughout lactation. To clarify the dose- response and kinetic characteristics of AsB and TMAO in seals, we conducted validation trials in which varying doses of biomarkers were fed to juvenile hooded seals (Cys- tophora cristata) in captivity at the University of Troms? in collaboration with E. S. Nord?y and A. S. Blix. Plasma TMAO peaked in about six to eight hours after intake and returned to low levels within 30 hours. Thus, biomarker methods may clarify whether a dive bout within a 24-hour period is associated with food capture, a considerable im- provement over arbitrary assignment of dive shapes to for- aging or nonforaging categories based on untested assump- tions about prey hunting behavior. ONTOGENY OF FORAGING IN WEDDELL SEAL PUPS A disconnect between maternal mass loss and pup mass gain could also arise because (1) pups begin to forage independently during the lactation period, leading to mass gain without corresponding maternal loss, or (2) variation in the pattern of energy deposition (e.g., fat versus protein) alters the pattern of pup mass gain independent of mater- nal expenditures. A prolonged period of dependence is characteristic of mammals in which development of social relationships appears to be important, such as in elephants, many pri- mates, and some odontocete whales (West et al., 2007). However, phocid seals typically wean their pups abruptly, with departure of the mother from the breeding colony. Foraging by suckling pups has so far been described for only two phocid species, the ringed seal Phoca hispida and the bearded seal Erignathus barbatus (Lydersen and Kovacs, 1999). Nursing Weddell seal pups commence div- ing at about two weeks of age and, on average, perform in excess of 20 dives per day (Burns and Testa, 1997). Al- though mothers and pups may at times dive together (Sato et al., 2002, 2003), it is unclear whether this entails any ?teaching? of the pup with regards to location, type, or capture of prey. There is a single published observation of the presence of milk and crustacean prey in the stom- ach of a Weddell seal pup (Lindsey, 1937), and pups have occasionally been observed bringing captured fi sh to the surface (K. Wheatley, University of Tasmania, personal communication, May 2004). It is possible that feeding by nursing pups could refl ect an inadequate rate of maternal energy transfer during lactation (e.g., Hayssen, 1993), but it is not known if feeding by pups is common or excep- tional in this species. UNCERTAINTIES ABOUT THE WEDDELL SEAL STRATEGY Much remains to be learned about the relative impor- tance of foraging (income) versus stored reserves (capital) in the lactation strategies of Weddell seals at both individ- ual and population levels. Weddell seals clearly rely exten- sively on stored reserves, but whether these are suffi cient to support the demands of lactation in some or all females is uncertain. Foraging is much more prevalent during the lactation period than previously thought, but the magni- tudes of energy and nutrient intakes are not known. It seems likely that access to food resources is important to reproductive success at least during the second half of lactation, when most females forage. In years when heavy multiyear ice has failed to break out of McMurdo Sound due to giant icebergs that have blocked egress to the north, the numbers of Weddell pups born has been reduced by up to 50%? 65% (R. Garrott, personal communication, 2007). By blocking light penetration the ice undoubtedly reduced 25_Eisert_pg335-346_Poles.indd 34225_Eisert_pg335-346_Poles.indd 342 11/17/08 9:18:20 AM11/17/08 9:18:20 AM WEDDELL SEAL CAPITAL BALANCE DURING LACTATION 343 primary productivity in McMurdo Sound, and this could result in a reduction in prey resources for Weddell seals. This may have led mothers to seek out alternative breed- ing sites in closer proximity to food resources, although the mechanism by which such choice is made is not known. It would be especially valuable to examine the varia- tion in foraging success and reproductive performance in different areas in the Antarctic where availability of food resources varies in time and space. How fl exible are the reproductive strategies of Weddell seals? Does the relative importance of income versus capital breeding vary among populations or among years? How could the reproductive success of the Weddell seal be impacted by changes in ice or currents associated with global warming? In a world of change, we need suffi cient background information on the resource needs of species to be able to predict future population trends. ACKNOWLEDGMENTS Weddell seal research at Hutton Cliffs, Erebus Bay, McMurdo Sound, Antarctica, in 1999 was supported by the Lottery Science Board New Zealand and the New Zea- land Antarctic Program. Research at this site in 2006 and 2007 was supported by the National Science Foundation Offi ce of Polar Programs (NSF-OPP award ANT-0538592) and authorized by permits issued by the National Marine Fisheries Service Offi ce of Protected Resources (permit 763-1485-00) and NSF-OPP (Antarctic Conservation Act permit 2007-001). We thank our fi eld teams for their hard work and dedication. LITERATURE CITED Agricultural Research Council. 1980. The Nutrient Requirements of Ruminant Livestock. Slough, U.K.: Commonwealth Agricultural Bureaux. Al-Waiz, M., S. C. Mitchell, J. R. Idle, and R. L. Smith. 1987. The Me- tabolism of 14 C-Labelled Trimethylamine and Its N-Oxide in Man. Xenobiotica, 17(5): 551? 558. Al-Waiz, M., M. Mikov, S. C. Mitchell, and R. L. Smith. 1992. 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Biological invasions in coastal ecosystems have occurred throughout Earth?s history, but the scale and tempo have increased greatly in recent time due to hu- man-mediated dispersal. Available data suggest that a strong latitudinal pattern exists for such human introductions in coastal systems. The documented number of introduced species (with established, self-sustaining populations) is greatest in temperate regions and declines sharply at higher latitudes. This observed invasion pattern across latitudes may result from differences in (1) historical baseline knowledge, (2) propagule supply, (3) resistance to invasion, and (4) disturbance regime. To date, the relative importance of these mechanisms across geographic regions has not been evaluated, and each may be expected to change over time. Of particular interest and concern are the interactive ef- fects of climate change and human activities on marine invasions at high latitudes. Shifts in invasion dynamics may be especially pronounced in the Northern Hemisphere, where current models predict not only an increase in sea surface temperatures but also a rapid reduction in sea ice in the Arctic. These environmental changes may greatly increase invasion opportunity at high northern latitudes due to shipping, mineral exploration, shoreline development, and other human responses. INTRODUCTION The extent and signifi cance of biological invasions in coastal marine ecosys- tems has become increasingly evident in recent years. On multiple continents, studies have described invasions by nonnative marine species, occurring pri- marily in shallow waters of bays and estuaries (e.g., Cohen and Carlton, 1995; Cranfi eld et al., 1998; Orensanz et al., 2002; Hewitt et al., 2004; Kerckhof et al., 2007; Fofonoff et al., in press). Although the ecological effects of most invasions have not been explored, it is evident that some nonnative species exert strong effects on the structure and function of invaded coastal ecosystems (Ruiz et al., 1999; Carlton, 2001; Grosholz, 2002). Marine invasions have occurred throughout Earth?s history, occurring some- times as punctuated events in geologic time that correspond to changes in cli- mate and dispersal barriers (e.g., Vermeij, 1991a, 1991b). However, invasions in modern time differ from those of the past, especially with respect to spatial and temporal scale. Most invasions are now driven primarily by the human-mediated transfer of organisms, instead of natural dispersal processes. As one consequence, Gregory M. Ruiz, Smithsonian Environmental Research Center, 647 Contees Wharf Road, Edge- water, MD 21037, USA. Chad L. Hewitt, Aus- tralian Maritime College, University of Tasmania, Bag 1370, Launceston, Tasmania 7250, Australia. Corresponding author: G. Ruiz (ruizg@si.edu). Accepted 28 May 2008. Latitudinal Patterns of Biological Invasions in Marine Ecosystems: A Polar Perspective Gregory M. Ruiz and Chad L. Hewitt 26_Ruiz_pg347-358_Poles.indd 34726_Ruiz_pg347-358_Poles.indd 347 11/17/08 9:17:49 AM11/17/08 9:17:49 AM 348 SMITHSONIAN AT THE POLES / RUIZ AND HEWITT the potential range of dispersal is arguably less constrained than in the past, without the need for geographic adjacency. Marine organisms are now often moved quickly by humans across great distances and dispersal barriers (e.g., ocean basins, hemispheres, and continents), which were previ- ously insurmountable in ecological timescales for most coastal species. Human transport of organisms also has increased the rate of invasions in recent time. It is clear that the docu- mented rate of marine invasions from human causes has increased dramatically, especially in the past 100? 200 years, in many global regions (e.g., Cohen and Carlton, 1998; Hewitt et al., 1999, 2004; Wonham and Carlton, 2005). The current tempo of invasions may, in fact, be unprecedented, resulting from the massive and growing scope of global trade, but it remains challenging to esti- mate actual rates of invasion that adequately control for potential biases (Ruiz et al., 2000). Nonetheless, a broad consensus exists that the pace of invasions has increased sharply in many well-studied regions. Despite considerable literature on patterns and pro- cesses of marine invasion, there is surprisingly little analysis of latitudinal patterns of invasion. In this paper, we review the current state of knowledge about human- mediated in- vasions (hereafter invasions or introductions) along a gra- dient from temperate to polar marine ecosystems, and we consider possible effects of climate change on invasions at high-latitudes. An extension of this comparison to tropical latitudes is the focus of future analyses. TEMPERATE-POLAR PATTERN OF INVASIONS For marine systems, most introductions (established, self-sustaining populations of nonnative species) are documented from temperate latitudes, including North America, Australia, Europe, New Zealand, and South America (see Cranfi eld et al., 1998; Hewitt et al., 1999, 2004; Reise et al., 1999; Ruiz et al., 2000; Hewitt, 2002; Orensanz et al., 2002; Castilla et al., 2005; Kerckhof et al., 2007; CIESM, 2007). While scores to hundreds of nonna- tive species are known from single bays and estuaries in temperate regions, few invasions are known from similar high-latitude sites, especially in polar regions. This pattern is illustrated by the sharp decline in docu- mented introductions with increasing latitude along west- ern North America (Figure 1). In an analysis of available literature and collection records, Ruiz et al. (2006a) exam- ined the number of nonnative marine invertebrate species reported from 12 large bays and estuaries (each including commercial ports) between 32? and 61?N latitude. For the six southernmost sites, from California to Washington, the number of documented introductions ranged from 50 to 170 species, with the largest number reported for San Francisco Bay (often the initial site for reported introduc- tions that spread to other bays; see Cohen and Carlton, 1995, for further discussion). In contrast, the six northern sites in Alaska were at least one order of magnitude lower in the number of known introductions, ranging from 0 to 5 species. For the entire Arctic (H1135066?N), we are aware of only one nonnative marine species that is known to have an established population. The Alaskan king crab, Paral- ithodes camtschaticus, was intentionally introduced to the White Sea in the 1960s to establish a fi shery and now occurs abundantly from Russia to Norway (Jorgensen, 2005). While other nonnative species have been reported for Arctic bioregions (e.g., Streftaris et al., 2005), it ap- pears that such occurrences either have not been docu- mented above 66?N or are not known to exist as estab- lished populations. For the Antarctic (H1102260?S), two nonnative species have been reported recently, both on the Antarctic Peninsula, but neither are known to have established populations. Two specimens of the North Atlantic spider crab Hyas araneus were found in collections from 1986, including one male and one female (Tavares and De Melo, 2004). In addition, the European green alga Ulva intestinalis was also reported (Clayton, 1997); however, the morphologi- cal identifi cation may be suspect. While these two species, FIGURE 1. Total number of nonnative marine invertebrate species with established populations in bays along western North America. Data are summarized from species reported in the literature and col- lections in each of 12 sites from California to Alaska (from Ruiz et al., 2006a). 26_Ruiz_pg347-358_Poles.indd 34826_Ruiz_pg347-358_Poles.indd 348 11/17/08 9:17:50 AM11/17/08 9:17:50 AM LATITUDINAL PATTERNS OF MARINE INVASIONS 349 and perhaps others, may have invaded the Antarctic, this has not been confi rmed to date (Lewis et al., 2003, 2004; Clarke et al., 2005). To some extent, the observed differences in nonnative species richness across latitudes may refl ect bias in search effort and taxonomic knowledge, which undoubtedly de- clines from temperate regions to the poles. It is virtually certain that other nonnative species are present at high lat- itudes and have not been recognized because of either lack of sampling or insuffi cient taxonomic and biogeographic resolution. However, such differences in historical baseline are unlikely to account for the overall latitudinal pattern, especially when considering the larger, conspicuous organ- isms (e.g., decapods, shelled molluscs, and ascidians). This is further supported by recent surveys in Alaskan waters that found a paucity of nonnative sessile invertebrates rel- ative to other sites in the continental United States (Ruiz et al., 2006a, unpublished data). The poleward decline in invasions apparently results from latitudinal differences in propagule supply of non- native species, resistance (or susceptibility) to invasion, or disturbance regimes. These may operate alone or in combination to produce the observed pattern of nonna- tive species richness. There exists theoretical and empirical support for the role of each factor in invasion dynamics (see Ruiz et al., 2000, and references therein), although these have not been evaluated for latitudinal patterns of marine invasions. Below, we consider each of these poten- tial mechanisms and how they may contribute to observed patterns in further detail, focusing particular attention on western North America. DIFFERENCES IN INVASION MECHANISMS ACROSS LATITUDES PROPAGULE SUPPLY The delivery pattern of organisms (propagules) greatly affects the likelihood of established populations. Propagule supply can be further divided into multiple components, including total number of propagules and the frequency (rate) and magnitude of inocula. Assuming suitable envi- ronmental conditions exist for a species to persist (including survival, growth, and successful reproduction), the likeli- hood of establishment is generally expected to increase with an increase in each component (Ruiz and Carlton, 2003; Lockwood et al., 2005; Johnston et al., in press). Most marine introductions are thought to result from species transfers by vessels and live trade. For North America, at least 50% of introduced marine species have been attributed to commercial ships, which move species associated with their underwater surfaces and also in bal- lasted materials (Ruiz et al., 2000; Fofonoff et al., 2003; see Carlton, 1985, for description of the history and use of solid ballast and ballast water). After shipping, live trade is the second largest mechanism (vector) of marine introduc- tions to North America, resulting from species transfers for aquaculture, fi sheries, bait, and aquaria (e.g., Cohen and Carlton, 1995; Carlton, 2001; Fofonoff et al., in press); in- vasions from live trade include both the target species of interest as well as many associated species, such as epibi- ota, parasites, and pathogens. These two vectors are active and often dominant throughout the world, although their relative importance certainly varies in space and time (e.g., Cranfi eld et al., 1998; Hewitt et al., 1999; 2004; Orensanz et al., 2002; Wasson et al., 2001; Castilla et al., 2005; see also Ribera and Boudouresque, 1995; Ribera Siguan, 2003; Hewitt et al., 2007). Once established, nonnative species often spread along the coast from the initial site of introduction. Some introduced marine species can expand their range in a new territory to encompass hundreds of kilometers (e.g., Grosholz, 1996; Thresher et al., 2005). This spread may occur by a combination of natural dispersal and anthro- pogenic means, depending upon the circumstances. Thus, invasion to a particular location can result by an initial introduction from distant sources or spread from an adja- cent population. In general, proximity to potential source populations may often increase the chances of coloniza- tion, especially for the latter. The current level of human activity, and especially shipping and live trade, is relatively low in polar regions, limiting opportunity for human-mediated transfers (e.g., Lewis et al., 2003, 2004). Moreover, the arrival of nonna- tive organisms from adjacent regions by natural dispersal is also likely to be low, resulting from a combination of low prevalence of nonnative species in adjacent regions and also the considerable distances or barriers that exist between potential sources for invasion of polar habitats. It is informative to compare the magnitude of commer- cial shipping to various regions of the United States (Fig- ure 2). For 2004? 2005, far fewer ship arrivals occurred in Alaska compared to other regions at lower latitudes. Unlike the latter regions, most ship arrivals to Alaska were from domestic sources, originating from other U.S. ports (par- ticularly those on the west coast) instead of foreign ports. Importantly, even the current level of shipping to Alaska is only a very recent development, increasing substantially over just the past few decades. Although these temporal changes in shipping have not been fully quantifi ed, an 26_Ruiz_pg347-358_Poles.indd 34926_Ruiz_pg347-358_Poles.indd 349 11/17/08 9:17:51 AM11/17/08 9:17:51 AM 350 SMITHSONIAN AT THE POLES / RUIZ AND HEWITT obvious increase has occurred. This is best exemplifi ed for oil tankers. Recent studies show a large number of marine organisms are delivered to Alaska in the ballast water of oil tankers. In 1998, it was estimated that oil tankers dis- charged a mean volume of 32,715 m 3 of ballast water per arrival to Port Valdez (61?N), containing an average density of 12,637 plankton per m 3 (as sampled by 80-H9262m mesh net, n H11005 169 vessels, chain-forming diatoms excluded; Hines and Ruiz, 2000). Most of these ships came from ports in California and Washington that are a potential source for many nonnative species (Figure 1). Over 17,000 oil tankers have arrived to Port Valdez since 1977, when the Alyeska pipeline was completed (Alyeska Pipeline Service Company, 2008). Prior to this date, tanker trade to Port Valdez simply did not exist. While we can consider the number of arrivals to be a coarse proxy for ship-mediated propagule supply to a re- gion, especially for a specifi c trade route and ship type (as above), this approach has clear limitations. Considerable variation exists among ships, voyage routes, and seasons in both the density and diversity of associated organisms (Smith et al., 1999; Coutts, 1999; Verling et al., 2005). In addition, the changing patterns of ship movements and trade present radically different invasion opportunities that are not captured in assessing the number of arrivals at one point in time (Hewitt et al., 1999, 2004; Minchin, 2006). As a result, the extent of species transfers by ships to locations is poorly resolved for any time period. Simi- lar limitations exist for most other transfer mechanisms in coastal ecosystems, making it challenging to estimate the actual propagule supply of nonnative species and provide direct comparisons across latitudes. Despite existing information gaps, the magnitude of nonnative propagule supply (both historically and pres- ently) has undoubtedly been low in polar regions. His- torically, whaling and sealing activities, particularly in the Southern Ocean (Murphy, 1995), provided some op- portunity for ship-mediated species transfer. Today, mod- ern shipping continues to provide a transfer mechanism to high latitudes in both hemispheres (Hines and Ruiz, 2000; Lewis, 2003, 2004). However, compared to tem- perate latitudes, the number of ship arrivals and the di- versity of routes (source ports) for the Arctic Ocean and Southern Ocean have been extremely limited. Rafting of marine species to the poles also appears to be low rela- tive to lower latitudes (Barnes, 2002). Finally, natural dis- persal of nonnative species is likely to be uncommon to both poles, perhaps especially in the Southern Hemisphere where distance and the Antarctic Circumpolar Circulation appear to create a signifi cant dispersal barrier (see reviews by Clarke et al., 2005, and Barnes et al., 2006). RESISTANCE TO INVASIONS Independent of propagule supply, high latitudes may be more resistant (less susceptible) to invasions. This can result from environmental resistance, whereby physical or chemical conditions in the recipient environment are not conducive to survivorship, reproduction, and population growth. Alternatively, biotic resistance can result from predators, competitors, food resources or other biological interactions that limit colonization success. There is support for environmental resistance to polar invasions due to the current temperature regime. In the Antarctic, low temperature is considered to be responsible through geologic time for the low diversity of decapod crustaceans, sharks, and other taxonomic groups and is also thought to operate today as a potential barrier to colonization (Thatje et al., 2005a, 2005b; Barnes et al., 2006; but see Lewis et al., 2003). This is, perhaps, best illustrated by research on lithodid crabs, which are physi- ologically unable to perform at the current polar tempera- tures (Aronson et al., 2007). In the Northern Hemisphere, using environmental niche models, deRivera et al. (2007) found that the north- ern ranges of some introduced species (the crab Carcinus maenas, the periwinkle Littorina saxatilis, the ascidian Styela clava, and the barnacle Amphibalanus improvisus) along western North America are not limited by tempera- ture. While none of these species appeared capable of FIGURE 2. Estimated number of commercial ship arrivals per year (2004? 2005) to regions of the United States. (Data are from Miller et al., 2007.) 26_Ruiz_pg347-358_Poles.indd 35026_Ruiz_pg347-358_Poles.indd 350 11/17/08 9:17:51 AM11/17/08 9:17:51 AM LATITUDINAL PATTERNS OF MARINE INVASIONS 351 colonizing the Arctic Ocean under current climatic con- ditions, their estimated ranges all included Alaskan wa- ters (where they do not presently occur). Thus, available analyses indicate both Antarctic and Arctic waters are currently beyond the thermal tolerance for some species, but the extent to which invasions can occur in these polar regions is not known. Temperature may often serve as a barrier to polar in- vasions by directly limiting larval development rate, which is highly temperature dependent (Thatje et al., 2003, 2005a, 2005b; deRivera et al., 2007). Many species are able to persist and grow at cold temperatures as juveniles or adults, compared to larvae that require warmer water for successful development to metamorphosis. High lati- tudes can also have short seasons when suffi cient food exists to sustain larval development. These direct and in- direct effects of temperature are thought to have greatly favored nonplanktotrophic larvae at high latitudes (see Thatje et al., 2005a, and references therein). Aside from food limitation, the potential importance of biotic resistance for high-latitude marine invasions is largely unexplored. Specifi cally, it is not clear whether competition or predation would greatly affect invasion establishment at high latitudes, especially in comparison to more temperate regions. As discussed by deRivera et al. (2007), it is conceiv- able that biotic interactions operate to reduce the likelihood of invasions to Alaska, despite suitable environmental con- ditions, but this hypothesis has not been tested empirically. More broadly, understanding biotic resistance to invasion is a critical gap in marine systems. DISTURBANCE REGIME Disturbance can play an important role in invasion dynamics through a variety of mechanisms. In general, the role of disturbance in invasion ecology has focused on changes in biotic interactions, thereby reducing biotic re- sistance to invasion. This can result from changes in com- petition that affect availability of resources, such as open space for colonization, food, or nutrients. Alternatively, changes in predation pressure can increase invasion op- portunities. While disturbance agents may operate directly to affect competition or predation, they can also have an indirect effect of these interactions. For example, changes in sediment or nutrient loading may cause a change in resi- dent community structure (including species composition or abundance) that affects the strength of biotic interac- tions for newly arriving species. In marine systems, literature on the role of distur- bance in invasions has focused primarily on anthropo- genic sources of disturbance (e.g., Ruiz et al., 1999; Piola and Johnston, 2006). Past studies have especially consid- ered effects of fi sheries exploitation, changes to habitat structure (whether physical or biological in nature), and chemical pollution. Invasions themselves have also been considered an important source of disturbance in cases where invasions either have (1) negative effects on resi- dent biota by reducing biotic resistance (Grosholz, 2005) or (2) positive effects on newly arriving species by provid- ing habitat, hosts, or food resources that were previously limiting (Simberloff and Von Holle, 1999). Anthropogenic disturbance can clearly play an impor- tant role in invasion dynamics, in terms of both establish- ment and abundance of nonnative species, for which there is strong theoretical and empirical underpinning. While considered likely to be a key factor in the high diversity of nonnative species in some estuaries, such as San Francisco Bay (Cohen and Carlton, 1998), the relative contribution of disturbance to observed invasion histories is not well understood, as there are many confounding variables that vary among sites (Ruiz et al., 1999). The magnitude of local and regional sources of an- thropogenic disturbance has been low in high-latitude and polar marine systems to date, compared to lower latitudes, refl ecting the low level of human activities. In contrast, climate change represents a global-scale and human-me- diated disturbance, with pronounced effects expected for high latitudes (Arctic Climate Impact Assessment, 2005; Intergovernmental Panel on Climate Change (IPCC), 2007). While several researchers have begun to explore the potential effects of climate change to marine invasions (Stachowicz et al., 2002; Occhipinti-Ambrogi, 2007), they have focused primarily on effects of temperature at mid- latitudes. Below, we expand this scope to examine poten- tial direct and indirect effects of projected temperature changes at high latitudes, considering especially the re- sponse of human activities and implications for changes in propagule supply, invasion resistance, and local/ regional disturbance. EFFECTS OF CLIMATE CHANGE ON INVASIONS Climate change is expected to affect many dimensions of coastal ecosystems, including temperature regimes, sea level, upwelling, ocean currents, storm frequency and mag- nitude, and precipitation patterns, which will infl uence land-sourced runoff, leading to changes in nutrients and tur- bidity (IPCC, 2007). Shifts in these key physical processes 26_Ruiz_pg347-358_Poles.indd 35126_Ruiz_pg347-358_Poles.indd 351 11/17/08 9:17:51 AM11/17/08 9:17:51 AM 352 SMITHSONIAN AT THE POLES / RUIZ AND HEWITT are expected to cause myriad and complex changes to the structure, dynamics, and function of biological commu- nities. The magnitude and rate of climate change are the focus of ongoing research, as are the expected changes to coastal ecosystems. One of the clear effects of climate change is elevated sea surface temperature, which is expected to be greatest at high latitudes and includes the complete loss of Arctic sea ice in the summer (IPCC, 2007). While models estimate the seasonal disappearance of Arctic sea ice in the next 50? 100 years, empirical measures suggest ice loss is occur- ring at a more rapid rate (Serreze et al., 2007; Stroeve et al., 2007, 2008). In contrast, no trend in sea ice change has been detected in the Antarctic (Cavalieri et al., 2003; but see Levebvre and Goosse, 2008). In Table 1, we consider some consequences of climate change in polar ecosystems for invasions, comparing potential responses in the Arctic and Antarctic. Our goal is to explore how the direct ef- fects of climate and indirect effects on human activities (in response to climatic shifts and associated opportunities) may affect invasion dynamics at high latitudes. Temperature is expected to have a direct effect on en- vironmental resistance to invasion. It is clear that thermal regime sets range limits of many species, and these species are expected to shift in response to changing temperatures (Stachowicz et al., 2002; Occhipinti-Ambrogi, 2007). The projected changes for the Arctic and Antarctic waters should allow species invasions (both natural and human- mediated) to occur that are not possible now due to con- straints in thermal tolerance and possibly food availability (e.g., Thatje et al., 2005a; deRivera et al., 2007; Aronson et al., 2007). Thus, we hypothesize increased invasions at both poles (Table 1), caused by natural dispersal and human-aided transport. However, the rate of new inva- sions may differ greatly between Arctic and Antarctic eco- systems, depending upon the interaction of temperature, propagule supply, and invasion resistance. Currents are expected to shift as a component of cli- mate change and will no doubt affect propagule supply. While considerable uncertainty exists about changes in ocean currents, we hypothesize (Table 1) that especially strong effects of currents on propagule supply may occur across the Arctic Ocean due to (1) loss of sea ice, poten- tially allowing greater water movement, and (2) the con- tinuous nature of the shoreline, providing adjacent source communities for transport. It appears likely that these ad- jacent communities will increase in species richness with rising temperature as many native and nonnative species expand their northern range limits in response. We posit that currents would therefore operate with temperature to facilitate species transport and coastwise spread across the Arctic. While currents can also supply larvae to the Ant- arctic (Barnes et al., 2006), we do not expect an analogous and increasing role with temperature rise there (Table 1), TABLE 1. Hypothesized effects of climate change on invasion dynamics in Arctic and Antarctic ecosystems. A plus (H11001) indicates projected increase in invasion resulting from specifi ed independent variables and response variables (mechanisms that affect invasion dynamics); a dash (? ) indicates no expected effect or ambiguous outcome. Projected changes for invasions Agent Independent variable(s) Response variable(s) Arctic Antarctic Climate Temperature Environmental resistance H11001 H11001 Currents Dispersal H11001 ? Commercial shipping Number of ship transits Dispersal, disturbance H11001 ? Shift in trade routes Dispersal H11001 Shoreline development Port development Dispersal, disturbance, new habitat H11001 ? Shoreline development Disturbance, new habitat H11001 Mineral extraction Dispersal, disturbance, new habitat H11001 ? Fisheries Natural stocks Dispersal, disturbance H11001 ? Aquaculture Dispersal, disturbance, new habitat H11001 Tourism Number of visits Dispersal, disturbance H11001 H11001 Debris Quantity of debris Dispersal, disturbance H11001 ? 26_Ruiz_pg347-358_Poles.indd 35226_Ruiz_pg347-358_Poles.indd 352 11/17/08 9:17:52 AM11/17/08 9:17:52 AM LATITUDINAL PATTERNS OF MARINE INVASIONS 353 due in part to the great distances from other coastal habi- tats and adjacent populations as well as the nature of the continuous Antarctic Circumpolar Circulation that cre- ates a signifi cant dispersal barrier. There is considerable historical precedent for trans- Arctic exchange of biota between the Pacifi c and the Atlan- tic. Vermeij (1991a) documented 295 species of molluscs that crossed the Arctic Ocean from the Pacifi c to the At- lantic (261 species) or in the reverse direction (34 species) after the opening of the Bering Strait in the early Pliocene. This analysis included both species that took part in the interchange and species that were descended from those that did. While the faunal exchange occurred in geologic time, it may have been punctuated and also underscores the potential for such current-mediated transport in the future. We are now exploring further the environmental conditions under which this interchange occurred to bet- ter understand its implication for expected climate change in the Arctic. It is also noteworthy that a diatom species of Pacifi c origin, Neodenticula seminae, was recently dis- covered in the North Atlantic, where it now appears to be well established (Reid et al., 2007). The species, detected from collections in 1998, is thought to be a recent trans- Arctic exchange, although the possibility of ship-mediated introduction via ballast water cannot be excluded. In response to projected temperature changes in polar regions, we expect major shifts in the level of many hu- man activities that will affect marine invasion dynamics at high latitudes and elsewhere. Moreover, we hypothesize that the human responses and effects on invasions will be asymmetrical, with the greatest changes in the Northern Hemisphere (Table 1). The greatest effects (and greatest asymmetries) are likely to result from increases in com- mercial shipping and shoreline development, followed by fi sheries and tourism. As sea ice disappears in the Arctic, the opportunities for shipping and mineral (especially oil) exploitation in- crease dramatically. If the Arctic becomes safe for naviga- tion, its use for commercial shipping would have strong economic incentive, reducing transit times and fuel costs. For example, the Northwest Passage between Europe and Asia is estimated to be approximately 9,000 km shorter than transiting the Panama Canal (Wilson et al., 2004). In addition, there is added expense (use fees) and often delays associated with the Panama Canal. Currently, 13,000? 14,000 vessels transit the Panama Canal per year (Ruiz et al., 2006b), and approximately 75% of these ves- sels are on routes in the Northern Hemisphere. Thus, a large number of vessels could benefi t from using an Arctic trade route, especially when considering additional ships now transiting the Suez Canal that may also save time and expense on this new route. A defl ection of shipping routes into the Arctic has several likely consequences from an invasion perspective. First, this would greatly increase the number of ships tran- siting the polar waters and thereby increase the delivery of nonnative propagules associated with hulls and ballast tanks to high latitudes. The per-ship magnitude of this supply is not immediately clear as it depends upon source. Second, the source ports for transiting vessels would in- crease the diversity of organisms being delivered. Because few ships now visit Arctic water, they certainly do not include the full selection of geographic source ports (and associated biotic assemblages) of ships now transiting the Panama and Suez canals. Third, ships also have chemi- cal discharges, whether intended or accidental (such as oil spills and leaching of active antifouling compounds), which represent a form of disturbance that may affect in- vasion resistance. On a broader geographic scale, we expect the de- crease in transit time may greatly improve survivorship of organisms associated with ballast water because survi- vorship in transit is time-dependent (Verling et al., 2005). The same is likely to be true for organisms on ships? hulls. Improved survivorship for either or both would result in increased propagule supply to subsequent ports of call, including major ports in Asia, Europe, and North America. While survivorship of shipborne organisms is time-dependent, the relative effects of transiting warm (Panama Canal) versus cold (Arctic) water will also af- fect the magnitude of change in mass fl ux of organisms among existing temperate ports. A cold temperature may serve to lower metabolic requirements, extending com- petency and survivorship of associated organisms. How- ever, the overall effects of ambient temperature are likely to be complex, varying with species and source regions, and remain to be explored. With warming temperatures and retreating sea ice, we expect a potentially large increase in shore-based activities in the Arctic, especially due to increase in mineral extrac- tion and export. We consider three different but related ac- tivities that can result in increased invasion opportunity. 1. Commercial port development will occur on some scale to support oil extraction offshore, where large re- serves exist. This will result in some increase in (1) propa- gule supply by ships, (2) local disturbance from chemical discharges from ship and port operations, and (3) local disturbance in the creation of novel habitat (rip rap, piers, etc.) associated with shoreline modifi cations. The latter is especially relevant given that many nonnative species are 26_Ruiz_pg347-358_Poles.indd 35326_Ruiz_pg347-358_Poles.indd 353 11/17/08 9:17:52 AM11/17/08 9:17:52 AM 354 SMITHSONIAN AT THE POLES / RUIZ AND HEWITT found on such artifi cial substrates (Cohen and Carlton, 1995; Hewitt et al., 2004; Glasby et al., 2007), which may be especially important focal areas for colonization. If local export of oil occurs by shipping, this could greatly increase the scale of port development as well as propa- gule supply, as exemplifi ed by oil export from Port Valdez (see above). 2. The scope of shoreline development, and especially associated habitat alteration and disturbance (as outlined above), is likely to exceed that for commercial shipping alone. Specifi cally, we expect some level of development to support oversight of territorial jurisdiction among Arctic countries, shore-based mineral extraction, tourism, and fi sheries. It is diffi cult to gauge the potential scale of such development, although it is noteworthy that several coun- tries have recently increased their presence in the Arctic (including military and surveying activities) in support of claims to Arctic territory and underlying mineral re- sources. 3. Offshore mineral extraction itself will also create opportunities for increased dispersal of propagules as well as some disturbance. It is not uncommon to use mobile drilling platforms for oil exploration, where the platforms are towed among sites at slow speeds. Although little studied, this movement can occur over great distances (i.e., across ocean basins) and may result in the transport organisms at much greater densities than found on op- erating ships because (1) the platforms sometimes reside at previous sites for long periods, accumulating dense as- semblages of organisms, and (2) the speed of transport is relatively slow, increasing the chance that organisms will remain associated. To our knowledge, strategies to assess or to reduce the associated risk of species transfers have not been explored for mobile drilling platforms. As with port development, offshore oil platforms, when fi xed, cre- ate artifi cial (novel) habitats and have some risk of chemi- cal discharge, and both types of disturbance may affect susceptibility to invasion. For the Antarctic, we do not expect commercial ship- ping or shoreline development to occur to any great ex- tent, simply because the same economic drivers do not ex- ist there at this point in time. There are not major shipping routes that would benefi t from transiting near Antarctica, and access to mineral and other resources is restricted un- der the Antarctic Treaty System. The potential exists for fi sheries to expand much more rapidly in the Arctic than in the Antarctic. This difference results in large part from access. The Arctic is in relatively close proximity to current centers of population and human activity, and a considerable history of fi sheries at northern high latitudes already exists in the Arctic. In contrast, ac- cess to fi sheries resources in Southern Ocean waters H1102260?S is managed under the Antarctic Treaty System, specifi cally the Convention on the Conservation of Antarctic Marine Living Resources. It is diffi cult to say whether aquaculture would occur to any extent in high-latitude systems; how- ever, current aquaculture trends indicate that it is highly likely. As discussed for commercial shipping, fi sheries ac- tivities can increase the levels of propagule supply and dis- turbance, with the latter resulting from operation of ships (and discharges), removing predators and competitors, and creating physical disturbance with fi shing gear (especially bottom trawls; Thrush et al., 1995). Tourism is already a growing industry to both the Arc- tic and Antarctic, and we expect this trend to continue. As with fi shing, the scope for growth appears greater in the Arctic, simply because of distance and access (including cost). As most tourism occurs by ships, the potential con- sequences for invasions are as outlined previously. Finally, we predict an increase in the quantity of hu- man-derived fl oating debris to occur in the Arctic and surrounding high latitudes in the Northern Hemisphere, coincident with increased levels of shipping, shoreline de- velopment, fi sheries, and tourism, as these are all potential sources for fl oating debris. While Barnes (2002) reported relatively few organisms colonizing fl oating debris at high latitudes, the number may also increase under warmer temperatures. Further, in the absence of sea ice in the sum- mer, the potential for longer-distance transport of fl oating material across the Arctic exists. Thus, we surmise that fl oating debris may play an important future role in the inoculation and, especially, regional spread of species in the Arctic, in contrast to much smaller changes expected in the Antarctic. CONCLUSIONS At the present time, very few introduced species are known from marine ecosystems at high latitudes in either hemisphere, especially for polar regions. This low number most likely results from a combination of low propagule supply of nonnative species and environmental resistance to invasion due to cold water temperatures and seasonal fl uctuations in resources. The relative lack of anthropogenic disturbance may also serve to limit invasion opportunity. With projected increases in temperature and the dis- appearance of Arctic sea ice in summer, we should expect invasions to increase as (1) temperatures fall within the thermal tolerance limits of organisms that are arriving 26_Ruiz_pg347-358_Poles.indd 35426_Ruiz_pg347-358_Poles.indd 354 11/17/08 9:17:52 AM11/17/08 9:17:52 AM LATITUDINAL PATTERNS OF MARINE INVASIONS 355 and (2) human-mediated responses to climate change in- crease the propagule supply and decrease the resistance to invasion (to the extent it exists) through disturbance. The change in invasion risk at high latitudes is expected to increase most in the Northern Hemisphere, driven by potentially large scale increases in the level of commercial shipping and shoreline development (especially associated with extraction of mineral resources). Fisheries, tourism, and fl oating debris also are likely to increase the opportu- nity for invasions, but to a much smaller degree. The consequences of climate change for invasions at high latitudes deserve serious attention from a conserva- tion and management perspective. While global shifts in climate (especially temperature) are underway and serve to increase chances of polar invasions, it appears that human responses to climate change will largely determine the number of invasions that occur. Although nonnative species can arrive to polar ecosystems by natural dispersal (Barnes et al., 2006), these regions are relatively isolated geographically, and the scope for human transport is far greater. Signifi cant efforts should now focus on under- standing and reducing the transfer of nonnative species to the poles, aiming to avoid the high number and sig- nifi cant impacts of introductions experienced in temperate waters. Efforts to minimize invasion risk at lower latitudes have employed several approaches that are applicable and should be adopted in polar regions. These are conceptually simple, focusing especially on (1) prevention or reduction of species transport by human activities and (2) detection of invasions by nonnative species. In many nations, regu- lations exist to greatly reduce the delivery of organisms that pose some risk of invasions (Ruiz and Carlton, 2003). While these sometimes focus on specifi c species that are known to survive or cause signifi cant impacts in a region, many strategies are now aimed at reducing transfers of all organisms associated with a known vector, especially be- cause the number of potential species is vast and the risks of colonization and impacts are simply not known (i.e., have not been examined) for most species. As an example, this approach of vector management is now being applied to commercial ships in many countries, where ships are required to treat their ballast water before discharging in coastal areas, reducing the concentrations of all coastal organisms (e.g., Minton et al., 2005). A comprehensive effort to reduce species transfers should include an assessment of potential human- mediated vectors as the basis for developing and implementing vec- tor management (Ruiz and Carlton, 2003). For polar sys- tems, policies could be adopted immediately to reduce transfers by commercial ships, extending efforts developed in temperate latitudes, since ships? ballast water and hulls are known to carry a risk of invasions (Ruiz et al., 2000; Fofonoff et al., 2003; Hewitt et al., 2004). Analyses of additional present and future vectors, such as oil drilling platforms and fi sheries activities, would be obvious next steps to estimate the potential magnitude of species trans- fers and to consider options for vector management. In addition to vector management, efforts to detect invasions and to measure temporal changes in invasions should be established in polar regions. Ideally, this would include an initial baseline survey and repeated surveys through time designed explicitly to test hypotheses about invasions (e.g., Ruiz and Hewitt, 2002) and address sev- eral management needs. First, these data would provide a measure of whether invasions are occurring and help identify specifi c vector(s) for management, providing feed- back on how well management strategies are working to limit invasions (Ruiz and Carlton, 2003). Second, result- ing detections of new invasions would also enable efforts to eradicate or control invasions, as deemed desirable. These strategies for prevention and detection are easily understood, but implementation is not so easy to achieve. Often, there are issues related to resources (time and fund- ing), limiting the desired scope of effort. In addition, there can be issues related to jurisdiction that further compli- cate implementation, resulting from political (geographic) boundaries and institutional (legal) authorities. Unfortu- nately, in temperate marine systems, these impediments to effective management strategies are often not overcome until a threshold of signifi cant ecological or economic im- pacts is reached. Since few invasions are known for polar systems to date, the opportunity now exists to implement manage- ment and policy that would greatly limit invasions and their unwanted impacts in these unique communities. Given the transboundary aspects of polar systems, there is a clear need for international cooperation and agreements in this area. We hope this article stimulates actions to eval- uate and reduce invasion risks at high latitudes, applying the principals, methods, and experiences from temperate marine systems around the globe. ACKNOWLEDGMENTS We wish to thank Michael Lang at the Smithsonian In- stitution Offi ce of the Under Secretary for Science for the opportunity to participate in the IPY Symposium. The con- tent of this manuscript benefi ted from discussions with Gail Ashton, Paul Fofonoff, Amy Freestone, Tuck Hines, Mark 26_Ruiz_pg347-358_Poles.indd 35526_Ruiz_pg347-358_Poles.indd 355 11/17/08 9:17:52 AM11/17/08 9:17:52 AM 356 SMITHSONIAN AT THE POLES / RUIZ AND HEWITT Minton, Whitman Miller, and Brian Steves, all of Smithso- nian Environmental Research Center; Catherine deRivera, University of Portland; and Gretchen Lambert. Research in Alaska was supported by National Sea Grant Program, Prince William Sound Citizens? Advisory Council, Smithso- nian Institution, and U.S. Fish and Wildlife Service. LITERATURE CITED Aleyeska Pipeline Service Company. About Us. http:// www .alyeska-pipe .com /about .html (accessed 2 May 2008). Arctic Climate Impact Assessment. 2005. Arctic Climate Impact Assess- ment. New York: Cambridge University Press. Aronson, R., S. Thatje, A. Clarke, L. Peck, D. Blake, C. Silga, and B. Seibel. 2007. 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Proceedings of the Royal Society of London, Series B, 272: 1249? 1256. 26_Ruiz_pg347-358_Poles.indd 35726_Ruiz_pg347-358_Poles.indd 357 11/17/08 9:17:53 AM11/17/08 9:17:53 AM 358 SMITHSONIAN AT THE POLES / RUIZ AND HEWITT Vermeij, G. J. 1991a. When Biotas Meet: Understanding Biotic Inter- change. Science, 253: 1099? 1104. ???. 1991b. Anatomy of an Invasion: The Trans-Arctic Interchange. Paleobiology, 17: 281? 307. Wasson, K., C. J. Zabin, L. Bedinger, M. C. Diaz, and J. S. Pearse. 2001. Biological Invasions of Estuaries without International Shipping: The Importance of Intraregional Transport. Biological Invasions, 102: 143? 153. Wilson, K. J., J. Falkingham, H. Melling, and R. De Abreu. 2004. ?Ship- ping in the Canadian Arctic.? In Proceedings of the IEEE Interna- tional Geoscience and Remote Sensing Symposium, 2004, IGARSS ?04. Volume 3, pp. 1853? 1856. Piscataway, N.J.: IEEE Press. Wonham, M., and J. T. Carlton. 2005. Cool-Temperate Marine Inva- sions at Local and Regional Scales: The Northeast Pacifi c Ocean as a Model System. Biological Invasions, 7(3): 369? 392. 26_Ruiz_pg347-358_Poles.indd 35826_Ruiz_pg347-358_Poles.indd 358 11/17/08 9:17:53 AM11/17/08 9:17:53 AM ABSTRACT. Four hundred thousand years after the Big Bang, electrons and nuclei com- bined to form atoms for the fi rst time, allowing a sea of photons to stream freely through a newly transparent universe. After billions of years, those photons, highly redshifted by the universal cosmic expansion, have become the cosmic microwave background (CMB) radiation we see coming from all directions today. Observation of the CMB is central to observational cosmology, and the Antarctic plateau is an exceptionally good site for this work. The fi rst attempt at CMB observations from the plateau was an expedition to the South Pole in December 1986 by the Radio Physics Research group at Bell Laboratories. No CMB anisotropies were observed, but sky noise and opacity were measured. The re- sults were suffi ciently encouraging that in the austral summer of 1988? 1989, three CMB groups participated in the ?Cucumber? campaign, where a temporary site dedicated to CMB anisotropy measurements was set up 2 km from South Pole Station. These were summer-only campaigns. Wintertime observations became possible in 1990 with the es- tablishment of the Center for Astrophysical Research in Antarctica (CARA), a National Science Foundation Science and Technology Center. The CARA developed year-round observing facilities in the ?Dark Sector,? a section of Amundsen? Scott South Pole Station dedicated to astronomical observations. The CARA scientists fi elded several astronomi- cal instruments: Antarctic Submillimeter Telescope and Remote Observatory (AST/RO), South Pole Infrared Explorer (SPIREX), White Dish, Python, Viper, Arcminute Cos- mology Bolometer Array Receiver (ACBAR), and Degree-Angular Scale Interferometer (DASI). By 2001, data from CARA, together with that from Balloon Observations of Millimetric Extragalactic Radiation and Geophysics (BOOMERANG? a CMB experi- ment on a long-duration balloon launched from McMurdo Station on the coast of Ant- arctica) showed clear evidence that the overall geometry of the universe is fl at, as opposed to being positively or negatively curved. In 2002, the DASI group reported the detection of polarization in the CMB. These observations strongly support a ?concordance model? of cosmology, where the dynamics of a fl at universe are dominated by forces exerted by the mysterious dark energy and dark matter. The CMB observations continue on the Ant- arctic plateau. The South Pole Telescope (SPT) is a newly operational 10-m-diameter off- set telescope designed to rapidly measure anisotropies on scales much smaller than 1?. INTRODUCTION Cosmology has made tremendous strides in the past decade; this is generally understood within the scientifi c community, but it is not generally appreciated that some of the most important results have come from Antarctica. Observational Robert W. Wilson and Antony A. Stark, Smithson- ian Astrophysical Observatory, 60 Garden Street, Cambridge, MA 02138, USA. Corresponding author: R. W. Wilson (rwilson@cfa.harvard.edu). Accepted 25 June 2008. Cosmology from Antarctica Robert W. Wilson and Antony A. Stark 27_Wilson_pg359-368_Poles.indd 35927_Wilson_pg359-368_Poles.indd 359 11/17/08 9:57:26 AM11/17/08 9:57:26 AM 360 SMITHSONIAN AT THE POLES / WILSON AND STARK cosmology has become a quantitative science. Cosmolo- gists describe the universe by a model with roughly a dozen parameters, for example, the Hubble constant, H 0 , and the density parameter, H9024. A decade ago, typical errors on these parameters were 30% or greater; now, most are known within 10%. We can honestly discriminate for and against cosmological hypotheses on the basis of quantita- tive data. The current concordance model, Lambda? Cold Dark Matter (Ostriker and Steinhardt, 1995), is both highly detailed and consistent with observations. This paper will review the contribution of Antarctic observa- tions to this great work. From the fi rst detection of the cosmic microwave back- ground (CMB) radiation (Penzias and Wilson, 1965), it was understood that deviations from perfect anisotropy would advance our understanding of cosmology (Peebles and Yu, 1970; Harrison, 1970): the small deviations from smoothness in the early universe are the seeds from which subsequent structure grows, and these small irregulari- ties appear as differences in the brightness of the CMB in various directions on the sky. When the universe was only 350,000 years old, the CMB radiation was released by elec- trons as they combined with nuclei into atoms for the fi rst time. Anisotropies in the CMB radiation indicate slight dif- ferences in density and temperature that eventually evolve into stars, galaxies, and clusters of galaxies. Observations at progressively higher sensitivity by many groups of sci- entists from the 1970s through the 1990s failed to detect the anisotropy (cf. the review by Lasenby et al., 1998). In the course of these experiments, observing techniques were developed, detector sensitivities were improved by orders of magnitude, and the effects of atmospheric noise became better understood. The techniques and detectors were so improved that the sensitivity of experiments came to be dominated by atmospheric noise at most observatory sites. Researchers moved their instruments to orbit, to balloons, and to high, dry observatory sites in the Andes and in Ant- arctica. Eventually, CMB anisotropies were detected by the Cosmic Background Explorer satellite (COBE; Fixsen et al., 1996). The ground-based experiments at remote sites also met with success. The spectrum of brightness in CMB vari- ations as a function of spatial frequency was measured by a series of ground-based and balloon-borne experiments, many of them located in the Antarctic. The data were then vastly improved upon by the Wilkinson Microwave An- isotropy Probe (WMAP; Spergel et al., 2003). The future Planck satellite mission (Tauber, 2005), expected to launch in 2008, will provide high signal-to-noise data on CMB anisotropy and polarization that will reduce the error on some cosmological parameters to the level of 1%. Even in the era of CMB satellites, ground-based CMB observations are still essential for reasons of fundamental physics. Cosmic microwave background radiation occurs only at wavelengths longer than 1 mm. The resolution of a telescope (in radians) is equal to the observed wavelength divided by the telescope diameter. To work properly, the overall accuracy of the telescope optics must be a small fraction of a wavelength. Observing the CMB at resolu- tions of a minute of arc or smaller therefore requires a telescope that is 10 m in diameter or larger, with an overall accuracy of 0.1 mm or better. There are no prospects for an orbital or airborne telescope of this size and accuracy in the foreseeable future. There is, however, important sci- ence to be done at high resolution, work that can only be done with a large ground-based telescope at the best pos- sible ground-based site? the Antarctic plateau. DEVELOPMENT OF ASTRONOMY IN THE ANTARCTIC Water vapor is the principal source of atmospheric noise in radio observations. Because it is exceptionally cold, the climate at the South Pole implies exception- ally dry observing conditions. As air becomes colder, the amount of water vapor it can hold is dramatically reduced. At 0?C, the freezing point of water, air can hold 83 times more water vapor than saturated air at the South Pole?s average annual temperature of H1100249?C (Goff and Gratch, 1946). Together with the relatively high altitude of the South Pole (2850 m), this means the water vapor content of the atmosphere above the South Pole is two or three orders of magnitude smaller than it is at most places on the Earth?s surface. This has long been known (Smythe and Jackson, 1977), but many years of hard work were needed to realize the potential in the form of new astro- nomical knowledge (cf. the recent review by Indermuehle et al., 2006). A French experiment, Emission Millimetrique (EMI- LIE) (Pajot et al., 1989), made the fi rst astronomical obser- vations of submillimeter waves from the South Pole during the austral summer of 1984? 1985. Emission Millimetrique was a ground-based, single-pixel bolometer dewar operat- ing at ?900H9262m and fed by a 45-cm off-axis mirror. It had successfully measured the diffuse galactic emission while operating on Mauna Kea in Hawaii in 1982, but the ac- curacy of the result had been limited by sky noise (Pajot et al., 1986). Martin A. Pomerantz, a cosmic ray researcher at Bartol Research Institute, encouraged the EMILIE group to relocate their experiment to the South Pole (Lynch, 1998). 27_Wilson_pg359-368_Poles.indd 36027_Wilson_pg359-368_Poles.indd 360 11/17/08 9:57:26 AM11/17/08 9:57:26 AM COSMOLOGY FROM ANTARCTICA 361 There they found better observing conditions and were able to make improved measurements of galactic emission. Pomerantz also enabled Mark Dragovan, then a re- searcher at Bell Laboratories, to attempt CMB anisotropy measurements from the pole. Dragovan et al. (1990) built a lightweight 1.2-m-offset telescope and were able to get it working at the pole with a single-pixel helium-4 bolo- meter during several weeks in January 1987 (see Figure 1). The results were suffi ciently encouraging that several CMB groups (Dragovan et al., 1989; Gaier et al., 1989; Meinhold et al., 1989; Peterson et al., 1989) participated in the ?Cu- cumber? campaign in the austral summer of 1988? 1989, where three Jamesway tents and a generator were set up at a temporary site dedicated to CMB anisotropy 2 km from South Pole Station in the direction of the international date line. These were summer-only campaigns, where instru- ments were shipped in, assembled, tested, used, disassem- bled, and shipped out in a single three-month-long sum- mer season. Considerable time and effort were expended in establishing and then demolishing observatory facilities, with little return in observing time. What little observing time was available occurred during the warmest and wet- test days of midsummer. Permanent, year-round facilities were needed. The Ant- arctic Submillimeter Telescope and Remote Observatory (AST/RO; Stark et al., 1997, 2001) was a 1.7-m- diameter offset Gregorian telescope mounted on a dedicated perma- nent observatory building. It was the fi rst radio telescope to operate year-round at the South Pole. The AST/RO was started in 1989 as an independent project, but in 1991 it be- came part of a newly founded National Science Foundation Science and Technology Center, the Center for Astrophysi- cal Research in Antarctica (CARA, http:// astro .uchicago .edu/ cara; cf. Landsberg, 1998). The CARA fi elded sev- eral telescopes: White Dish (Tucker et al., 1993), Python ( Dragovan et al., 1994; Alvarez, 1995; Ruhl et al., 1995; Platt et al., 1997; Coble et al., 1999), Viper (Peterson et al., 2000), the Degree-Angular Scale Interferometer (DASI; Leitch et al., 2002a), and the South Pole Infrared Explorer (SPIREX; Nguyen et al., 1996), a 60-cm telescope operating primarily in the near-infrared K band. These facilities were housed in the ?Dark Sector,? a grouping of buildings that includes the AST/RO building, the Martin A. Pomerantz Observatory building (MAPO), and a new ?Dark Sector Laboratory? (DSL), all located 1 km away from the main base across the aircraft runway in a radio quiet zone. The combination of White Dish, Python, and Uni- versity of California at Santa Barbara 1994 (Ganga et al., 1997) data gave the fi rst indication, by 1997, that the spectrum of spatial anisotropy in the CMB was consistent with a fl at cosmology. Figure 2 shows the state of CMB anisotropy measurements as of May 1999. The early South Pole experiments, shown in green, clearly delineate a peak in CMB anisotropy at a scale H5129 H11005 200, or 1?, consistent with a fl at H9024 0 H11005 1 universe. Shortly thereafter, the Balloon Observations of Millimetric Extragalactic Radiation and Geophysics (BOOMERANG)-98 long-duration balloon experiment (de Bernardis et al., 2000; Masi et al., 2006, 2007; Piacentini et al., 2007) and the fi rst year of DASI (Leitch et al., 2002b) provided signifi cantly higher signal- to-noise data, yielding H9024 0 H11005 1 with errors less than 5%. This was a stunning achievement, defi nitive observations of a fl at universe balanced between open and closed Fried- mann solutions. In its second year, a modifi ed DASI made the fi rst measurement of polarization in the CMB (Kovac et al., 2002; Leitch et al., 2002c). The observed relationship between polarization and anisotropy amplitude provided a detailed confi rmation of the acoustic oscillation model of CMB anisotropy (Hu and White, 1997) and strong support for the standard model. The demonstration that the geom- etry of the universe is fl at is an Antarctic result. SITE TESTING One of the primary tasks for the CARA collaboration was the characterization of the South Pole as an observa- tory site (Lane, 1998). It proved unique among observatory sites for unusually low wind speeds, the complete absence of rain, and the consistent clarity of the submillimeter sky. FIGURE 1. Mark Dragovan, Robert Pernic, Martin Pomerantz, Robert Pfeiffer, and Tony Stark with the AT&T Bell Laboratories 1.2-m horn antenna at the South Pole in January 1987. This was the fi rst attempt at a CMB measurement from the South Pole. 27_Wilson_pg359-368_Poles.indd 36127_Wilson_pg359-368_Poles.indd 361 11/17/08 9:57:26 AM11/17/08 9:57:26 AM 362 SMITHSONIAN AT THE POLES / WILSON AND STARK Schwerdtfeger (1984) and Warren (1996) have compre- hensively reviewed the climate of the Antarctic plateau and the records of the South Pole meteorology offi ce. Chamberlin (2002) analyzed weather data to determine the precipitable water vapor (PWV), a measure of total water vapor content in a vertical column through the at- mosphere. He found median wintertime PWV values of 0.3 mm over a 37-year period, with little annual variation. The PWV values at South Pole are small, stable, and well- understood. Submillimeter-wave atmospheric opacity at South Pole has been measured using sky dip techniques. Chamberlin et al. (1997) made over 1,100 sky dip observations at 492 GHz (?609H9262m) with AST/RO during the 1995 observing season. Even though this frequency is near a strong oxy- gen line, the opacity was below 0.70 half of the time dur- ing the austral winter and reached values as low as 0.34, better than ever measured at any other ground-based site. From early 1998, the ?350H9262m band has been continuously monitored at Mauna Kea, Chajnantor, and the South Pole by identical tipper instruments developed by S. Radford of the National Radio Astronomy Observatory and J. Peter- son of Carnegie Mellon University and CARA. The 350- H9262m opacity at the South Pole is consistently better than at Mauna Kea or Chajnantor. Sky noise is caused by fl uctuations in total power or phase of a detector caused by variations in atmospheric emissivity and path length on timescales of order one second. Sky noise causes systematic errors in the mea- surement of astronomical sources. This is especially im- portant at the millimeter wavelengths for observations of the CMB: at millimeter wavelengths, the opacity of the atmosphere is at most a few percent, and the contribution to the receiver noise is at most a few tens of degrees, but sky noise may still set limits on observational sensitiv- ity. Lay and Halverson (2000) show analytically how sky noise causes observational techniques to fail: fl uctuations in a component of the data due to sky noise integrate down more slowly than t H110021/2 and will come to dominate the error during long observations. Sky noise at South Pole is considerably smaller than at other sites, even com- paring conditions of the same opacity. The PWV at the South Pole is often so low that the opacity is dominated by the dry-air component (Chamberlin and Bally, 1995; Chamberlin, 2002); the dry-air emissivity and phase error do not vary as strongly or rapidly as the emissivity and phase error due to water vapor. Lay and Halverson (2000) compared the Python experiment at the South Pole ( Dragovan et al., 1994; Alvarez, 1995; Ruhl et al., 1995; Platt et al., 1997; Coble et al., 1999) with the Site Test Interferometer at Chajnantor (Holdaway et al., 1995; Radford et al., 1996) and found that the amplitude of the sky noise at the South Pole is 10 to 50 times less than that at Chajnantor (Bussmann et al., 2004). The best observing conditions occur only at high el- evation angles, and at the South Pole this means that only the southernmost 3 steradians of the celestial sphere are accessible with the South Pole?s uniquely low sky noise, but this portion of sky includes millions of galaxies and cosmological sources, the Magellanic clouds, and most of the fourth quadrant of the galaxy. The strength of the South Pole as a millimeter and submillimeter site lies in the low sky noise levels routinely obtainable for sources around the south celestial pole. 10 1 10 2 10 3 10 4 May 1999 ? WD, ACTA, VLA CAT OVRO MAX MSAM COBE BAM Tenerife Saskatoon Viper SP 98 Python V SP 96 Python SP 91? 95 UCSB SP 94 0 Planck SP 10m H9024 0 H110051.0 0.1 20 40 60 80 100 FIGURE 2. Microwave background anisotropy measurements as of May 1999, prior to the launch of BOOMERANG, the deploy- ment of DASI , and the launch of WMAP. South Pole experimen- tal results are shown in green. Note that the peak at H5129 H11005 200 is clearly defi ned, indicating a fl at universe (H9024 0 H11005 1). Abbreviations are as follows: UCSB SP 94 H11005 a campaign at the South Pole in 1994 by the University of California at Santa Barbara sponsored by NSF; BAM H11005 Balloon-borne Anisotropy Measurement; COBE H11005 Cosmic Background Explorer; MSAM H11005 Medium Scale Anisotropy Mea- surement experiment; MAX H11005 Millimeter Anisotropy eXperiment; CAT H11005 Cosmic Anisotropy Telescope; OVRO H11005 Owens Valley Ra- dio Observatory; WD H11005 White Dish; ACTA H11005 Australia Telescope Compact Array; VLA H11005 Very Large Array. 27_Wilson_pg359-368_Poles.indd 36227_Wilson_pg359-368_Poles.indd 362 11/17/08 9:57:28 AM11/17/08 9:57:28 AM COSMOLOGY FROM ANTARCTICA 363 TELESCOPES AND INSTRUMENTS AT THE SOUTH POLE Viper was a 2.1-m off-axis telescope designed to allow measurements of low-contrast millimeter-wave sources. It was mounted on a tower at the opposite end of MAPO from DASI. Viper was used with a variety of instruments: Dos Equis, a CMB polarization receiver operating at 7 mm; the Submillimeter Polarimeter for Antarctic Re- mote Observing (SPARO), a bolometric array polarim- eter operating at ?450H9262m; and the Arcminute Cosmology Bolometer Array Receiver (ACBAR), a multiwavelength bolometer array used to map the CMB. The ACBAR is a 16-element bolometer array operating at 300 mK. It was specifi cally designed for observations of CMB anisotropy and the Sunyaev? Zel?dovich effect (SZE). It was installed on the Viper telescope early in 2001 and was successfully operated until 2005. The ACBAR has made high-quality maps of SZE in several nearby clusters of galaxies and has made signifi cant measurements of anisotropy on the scale of degrees to arcminutes (Runyan et al., 2003; Reichardt et al., 2008). The Submillimeter Polarimeter for Antarctic Remote Observing (SPARO) was a nine-pixel polarimetric imager operating at ?450H9262m. It was operational on the Viper tele- scope during the early austral winter of 2000. Novak et al. (2000) mapped the polarization of a region of the sky (H110110.25 square degrees) centered approximately on the Ga- lactic Center. Their results imply that within the Galactic Center molecular gas complex, the toroidal component of the magnetic fi eld is dominant. The data show that all of the existing observations of large-scale magnetic fi elds in the Galactic Center are basically consistent with the ?magnetic outfl ow? model of Uchida et al. (1985). This magnetody- namic model was developed in order to explain the Galactic Center radio lobe, a limb-brightened radio structure that extends up to one degree above the plane and may represent a gas outfl ow from the Galactic Center. The Degree Angular Scale Interferometer (DASI; Leitch et al., 2002a) was a compact centimeter-wave inter- ferometer designed to image the CMB primary anisotropy and measure its angular power spectrum and polariza- tion at angular scales ranging from two degrees to several arcminutes. As an interferometer, DASI measured CMB power by simultaneous differencing on several scales, measuring the CMB power spectrum directly. The DASI was installed on a tower adjacent to MAPO during the 1999? 2000 austral summer and had four successful win- ter seasons. In its fi rst season, DASI made measurements of CMB anisotropy that confi rmed with high accuracy the ?concordance? cosmological model, which has a fl at geometry, and made signifi cant contributions to the total stress energy from dark matter and dark energy (Halverson et al., 2002; Pryke et al., 2002). In its second year, DASI made the fi rst measurements of ?E-mode? polarization of the CMB (Leitch et al., 2002c; Kovac et al., 2002). The Antarctic Submillimeter Telescope and Remote Observatory (AST/RO) was a general-purpose 1.7-m- diameter telescope (Stark et al., 1997, 2001) for astron- omy and aeronomy studies at wavelengths between 200 and 2,000 H9262m. It was operational from 1995 through 2005 and was located in the Dark Sector on its own building. It was used primarily for spectroscopic studies of neutral atomic carbon and carbon monoxide in the in- terstellar medium of the Milky Way and the Magellanic Clouds. Six heterodyne receivers and a bolometer array were used on AST/RO: (1) a 230-GHz superconduc- tor- insulator-superconductor (SIS) receiver (Kooi et al., 1992), (2) a 450- to 495-GHz SIS quasi-optical receiver (Zmuidzinas and LeDuc, 1992; Engargiola et al., 1994), (3) a 450- to 495-GHz SIS waveguide receiver (Walker et al., 1992; Kooi et al., 1995), which could be used simulta- neously with (4) a 800- to 820-GHz fi xed-tuned SIS wave- guide mixer receiver (Honingh et al., 1997), (5) the Pole Star array, which deployed four 800- to 820-GHz fi xed- tuned SIS waveguide mixer receivers (see http:// soral .as .arizona .edu/ pole-star; Groppi et al., 2000; Walker et al., 2001), (6) the Terahertz Receiver with NbN HEB Device (TREND), a 1.5-THz heterodyne receiver ( Gerecht et al., 1999; Yngvesson et al., 2001), and (7) the South Pole Imaging Fabry-Perot Interferometer (SPIFI; Swain et al., 1998). Spectral lines observed with AST/RO included CO J H11005 7 j 6, CO J H11005 4 j 3, CO J H11005 2 j 1, HDO J H11005 1 0,1 j 0 0,0 , [C I] 3 P 1 j 3 P 0 , [C I] 3 P 2 j 3 P 1 , and [ 13 C I] 3 P 2 j 3 P 1 . There were four acousto- optical spectro meters (AOS; Schieder et al., 1989): two low- resolution spectrometers with a bandwidth of 1 GHz, an array AOS with four low- resolution spectrometer channels with a bandwidth of 1 GHz for the PoleSTAR array, and one high-resolution AOS with 60-MHz bandwidth. The Antarctic Submilli- meter Telescope and Remote Observatory produced data for over a hundred scientifi c papers relating to star forma- tion in the Milky Way and the Magellanic Clouds. Among the more signifi cant contributions is a submillimeter-wave spectral line survey of the Galactic Center region (Martin et al., 2004) that showed the episodic nature of starburst and black hole activity in the center of our galaxy (Stark et al., 2004). 27_Wilson_pg359-368_Poles.indd 36327_Wilson_pg359-368_Poles.indd 363 11/17/08 9:57:29 AM11/17/08 9:57:29 AM 364 SMITHSONIAN AT THE POLES / WILSON AND STARK The Q and U Extra-Galactic Sub-mm Telescope (QUEST) at DASI (QUaD; Church et al., 2003) is a CMB polarization experiment that placed a highly symmetric antenna feeding a bolometer array on the former DASI mount at MAPO, becoming operational in 2005. It is capable of measuring amplitude and polarization of the CMB on angular scales as small as 0.07?. The QUaD has suffi cient sensitivity to detect the conversion of E-mode CMB polarization to B-mode polarization caused by grav- itational lensing in concentrations of dark matter. Background Imaging of Cosmic Extragalactic Polar- ization (BICEP) (Keating et al., 2003; Yoon et al., 2006) is a millimeter-wave receiver designed to measure polariza- tion and amplitude of the CMB over a 20? fi eld of view with 1? resolution. It is mounted on the roof of the Dark Sector Laboratory and has been operational since early 2006. The design of BICEP is optimized to eliminate sys- tematic background effects and thereby achieve suffi cient polarization sensitivity to detect the component of CMB polarization caused by primordial gravitational waves. These measurements test the hypothesis of infl ation during the fi rst fraction of a second after the Big Bang. The South Pole Telescope is a 10-m-diameter off-axis telescope that was installed during the 2006? 2007 season (Ruhl et al., 2004). It is equipped with a large fi eld of view (Stark, 2000) that feeds a state-of-the-art 936-element bo- lometer array receiver. The initial science goal is a large SZE survey covering 4,000 square degrees at 1.3H11032 resolu- tion with 10 H9262K sensitivity at a wavelength of 2 mm. This survey will fi nd all galaxy clusters above a mass limit of 3.5 H11003 10 14 M H17018 , regardless of redshift. It is expected that an unbiased sample of approximately 8,000 clusters will be found, with over 300 at redshifts greater than one. The sample will provide suffi cient statistics to use the density of clusters to determine the equation of state of the dark energy component of the universe as a function of time. CONCLUSIONS Observations from the Antarctic have brought re- markable advances in cosmology. Antarctic observations have defi nitively demonstrated that the geometry of the universe is fl at. These observations were made possible by excellent logistical support offered for the pursuit of science at the Antarctic bases. 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IEEE Transactions on Microwave Theory and Techniques, 40: 1797. 27_Wilson_pg359-368_Poles.indd 36727_Wilson_pg359-368_Poles.indd 367 11/17/08 9:57:32 AM11/17/08 9:57:32 AM 27_Wilson_pg359-368_Poles.indd 36827_Wilson_pg359-368_Poles.indd 368 11/17/08 9:57:32 AM11/17/08 9:57:32 AM ABSTRACT. Feeling a bit hungry? Imagine that you only received one meal every few million years and that when you ate it, it was a gigantic Thanksgiving feast. That sort of gorging might give you quite a stomach ache! The black hole at the center of our galaxy seems to go through just this cycle of feast and famine, but as the turkey dinner arrives, it bursts into a tremendous display of fi reworks. Instead of turkey, a black hole eats a vast platter of dust and gas that is compressed and stressed as it reaches the inner part of the galaxy. This compression causes the formation of a plethora of large short-lived stars that go supernova shortly after their birth. These supernova fi reworks would then be suf- fi ciently intense to make the center of the galaxy one of the brightest objects in our night sky while at the same time sterilizing any life that might be nearby. How does this matter get to the center of the galaxy and when can we expect the next burst of fi reworks? At this very moment the dinner plate for the black hole at the center of the Milky Way is being assembled, and a group of astronomers from the South Pole is looking at the menu. Dinner will be served in about 10 million years. ANTARCTIC SUBMILLIMETER TELESCOPE AND REMOTE OBSERVATORY Observing the gas in the center of our galaxy requires an instrument that is sensitive to its emissions. While the bulk of the baryonic matter in our universe is simply hydrogen, it is surprisingly diffi cult to observe. Rather than detecting the hydrogen directly, tracers such as carbon monoxide (CO) are used instead to determine the density, temperature, and dynamics of the hydrogen with which it is well mixed. The Antarctic Submillimeter Telescope and Remote Observa- tory (AST/RO, Figure 1), located at 2847 m altitude at the Amundsen-Scott South Pole Station, was built in 1995 to study the emissions of these tracers. One might reasonably ask, why would one put a telescope like this at the bot- tom of the Earth? The South Pole has very low water vapor, high atmospheric stability, and a thin troposphere, making it exceptionally good for submillimeter observations (Chamberlin et al., 1997; Lane, 1998). Technically, AST/RO is a 1.7-m-diameter, offset Gregorian telescope capable of observing at wavelengths between 200 H9262m and 1.3 mm (Stark et al., 2001). The observations of the center of the Milky Way described here were taken during the austral winter seasons of 2001? 2004 by the telescope?s dedicated winter-over staff. Christopher L. Martin, Oberlin College, Depart- ment of Physics and Astronomy, 110 North Pro- fessor Street, Oberlin, OH 44074, USA (chris. martin@oberlin.edu). Accepted 25 June 2008. Feeding the Black Hole at the Center of the Milky Way: AST/RO Observations Christopher L. Martin 28_Martin_pg369-372_Poles.indd 36928_Martin_pg369-372_Poles.indd 369 11/17/08 9:40:52 AM11/17/08 9:40:52 AM 370 SMITHSONIAN AT THE POLES / MARTIN WHAT DOES THE GALACTIC CENTER LOOK LIKE WHEN SEEN BY AST/RO? Figure 2 presents spatial? spatial (l, b) maps integrated over velocity for the three transitions AST/RO has ob- served in the Galactic Center. The fi rst thing you might notice when looking at the maps is that the Galactic Cen- ter appears to be made up of clouds rather than stars. This is because the frequencies of light observed by AST/RO are sensitive to the dust and gas between the stars rather than the stars themselves. Looking beyond this, the next most striking result is that CO J H11005 4 j 3 emission in the Galactic Center region is essentially coextensive with the emission from the lower J transitions of CO. This con- trasts sharply with the outer galaxy where CO J H11005 4 j 3 emission is rather less extensive than CO J H11005 1j 0. On the other hand, CO J H11005 7 j 6 emission is very compact and constrained to just a few key regions, indicating high densities and temperatures in those regions. Four major cloud complexes are seen in the maps, from left to right: the complex at l H11015 1.3?, the Sgr B complex near l H11015 0.7?, the Sgr A cloud near l H11015 0.0?, and the Sgr C cloud near (l H11015 H110020.45?, b H11015 H110020.2?). As noted by Kim et al. (2002), the CO J H11005 7 j 6 emission is much more spatially con- fi ned than the lower-J CO transitions. In contrast, the [CI] (atomic carbon) emission is comparable in spatial extent to the low-J CO emission, but its distribution appears somewhat more diffuse (less peaked). The Sgr C cloud is much less prominent in the [CI] map than in the other fi ve transitions (Ojha et al., 2001). In order to understand what this gas is doing, we need to combine the data taken with AST/RO with data from other sources, and we need a broader understanding of the dynamics of the Galactic Center. FIGURE 1. The Antarctic Submillimeter Telescope and Remote Observatory (AST/RO) as it appeared shortly after its construction in 1995 with the main Amundsen-Scott South Pole Station in the background. 28_Martin_pg369-372_Poles.indd 37028_Martin_pg369-372_Poles.indd 370 11/17/08 9:40:53 AM11/17/08 9:40:53 AM AST/RO BLACK HOLE OBSERVATIONS 371 WHAT IS THIS GAS DOING? Much has been learned about dense gas in the Galactic Center region through radio spectroscopy. Early observa- tions of F(2 j 2) OH absorption (Robinson et al., 1964; Goldstein et al., 1964) suggested the existence of copious molecular material within 500 pc of the Galactic Center. This was confi rmed by detection of extensive J H11005 1 j 0 12 CO emission (Bania, 1977; Liszt and Burton, 1978). Subsequent CO surveys (Bitran, 1987; Stark et al., 1988; Bitran et al., 1997; Oka et al., 1998) have measured this emission with improving coverage and resolution. These surveys show a complex distribution of emission, which is chaotic, asymmetric, and nonplanar; there are hundreds of clouds, shells, arcs, rings, and fi laments. On scales of 100 pc to 4 kpc, however, the gas is loosely organized around closed orbits in the rotating potential of the underlying stellar bar (Binney et al., 1991). Some CO-emitting gas is bound into clouds and cloud complexes, and some is sheared by tidal forces into a molecular intercloud medium of a kind not seen elsewhere in the galaxy (Stark et al., 1989). The large cloud complexes, Sgr A, Sgr B, and Sgr C, are the among the largest molecular cloud complexes in the galaxy (M H11022 10 6.5 M sun ). Such massive clouds must be sinking toward the center of the galactic gravitational well as a result of dynamical friction and hydrodynamic effects (Stark et al., 1991). The deposition of these massive lumps of gas upon the center could fuel a starburst or an eruption of the central black hole (Genzel and Townes, 1987; Stark et al., 2004). To better understand the molecular gas of the Galactic Center, we need to determine its physical state? its tem- perature and density. This involves understanding radia- tive transfer in CO, the primary tracer of molecular gas. Also useful would be an understanding of the atomic car- bon lines, [CI], since those lines trace the more diffuse mo- lecular regions, where CO is destroyed by UV radiation but H 2 is still present. Hence, a key project of the Antarctic Submillimeter Telescope and Remote Observatory reported by Martin et al. (2004) has been the mapping of CO 4? 3 and CO 7? 6 emission from the inner Milky Way, allowing deter- mination of gas density and temperature. Galactic Center gas that Binney et al. (1991) identify as being on x 2 orbits has a density near 10 3.5 cm 3 , which renders it only mar- ginally stable against gravitational coagulation into a few giant molecular clouds. This suggests a relaxation oscilla- tor mechanism for starbursts in the Milky Way, whereby infl owing gas accumulates in a ring at 150-pc radius until the critical density is reached and the resulting instability leads to the sudden formation of giant clouds and the de- position of 4 H11003 10 7 M sun , of gas onto the Galactic Center. Depending on the accretion rate near the inner Lindblad resonance, this cycle will repeat with a timescale on the order of 20 Myr, leading to starbursts on the same time- scale. When we analyze our data (Stark et al., 2004), we FIGURE 2. Spatial? spatial (l, b) integrated intensity maps for the three transitions observed with AST/RO. Transitions are identifi ed at the left on each panel. The emission is integrated over all velocities where data are available. All three maps have been smoothed to the same 2H11032 resolu- tion. Electronic versions of results from this region as published in Martin et al. (2004) may be requested from the author via e-mail. 28_Martin_pg369-372_Poles.indd 37128_Martin_pg369-372_Poles.indd 371 11/17/08 9:40:57 AM11/17/08 9:40:57 AM 372 SMITHSONIAN AT THE POLES / MARTIN observe gas approaching this density and thus beginning the next stage of its long descent in the black hole at the heart of our galaxy. In the Thanksgiving feast analogy, the gas that we can see coagulating on these outer orbits and beginning its slow trip inward is the future dinner for the black hole at the center of our galaxy. When it arrives at the center of the galaxy, it will provide the fuel for a burst of star formation activity (a starburst) at the very heart of the Milky Way and, hence, for the fi reworks that will light up our night skies in the millennia to come. ACKNOWLEDGMENTS This research was supported in part by the National Science Foundation under NSF grant number ANT- 0126090 and was conducted with the assistance of the many collaborators of the Antarctic Submillimeter Tele- scope Remote Observatory (AST/RO). LITERATURE CITED Bania, T. M. 1977. Carbon Monoxide in the Inner Galaxy. Astrophysical Journal, 216: 381? 403. Binney, J., O. E. Gerhard, A. A. Stark, J. Bally, and K. I. Uchida. 1991. Understanding the Kinematics of Galactic Centre Gas. Monthly Notices of the Royal Astronomical Society, 252: 210? 218. Bitran, M., H. Alvarez, L. Bronfman, J. May, and P. Thaddeus. 1997. A Large Scale CO Survey of the Galactic Center Region. Astronomy and Astrophysics, Supplement Series, 125: 99? 138. Bitran, M. E. 1987. CO in the Galactic Center: A Complete Survey of CO Emission in the Inner 4 kpc of the Galaxy. Ph.D. diss., Univer- sity of Florida, Gainesville. Chamberlin, R. A., A. P. Lane, and A. A. Stark. 1997. The 492 GHz Atmospheric Opacity at the Geographic South Pole. Astrophysical Journal, 476: 428? 433. Genzel, R., and C. H. Townes. 1987. Physical Conditions, Dynamics, and Mass Distribution in the Center of the Galaxy. Annual Review of Astronomy and Astrophysics, 25: 377? 423. Goldstein, S. J., E. J. Gundermann, A. A. Penzias, and A. E. Lilley. 1964. OH Absorption Spectra in Sagittarius. Nature, 203: 65? 66. Kim, S., C. L. Martin, A. A. Stark, and A. P. Lane. 2002. AST/RO Obser- vations of CO J H11005 7j6 and J H11005 4j3 Emission toward the Galactic Center Region. Astrophysical Journal, 580: 896? 903. Lane, A. P. 1998. ?Submillimeter Transmission at South Pole.? In As- trophysics From Antarctica, ed. G. Novack and R. H. Landsberg, p. 289. Astronomical Society of the Pacifi c Conference Series, No. 141. San Francisco: Astronomical Society of the Pacifi c. Liszt, H. S., and W. B. Burton. 1978. The Gas Distribution in the Central Region of the Galaxy. II? Carbon Monoxide. Astrophysical Jour- nal, 226: 790? 816. Martin, C. L., W. M. Walsh, K. Xiao, A. P. Lane, C. K. Walker, and A. A. Stark. 2004. The AST/RO Survey of the Galactic Center Region I. The Inner 3 Degrees. Astrophysical Journal, Supplement Series, 150: 239? 262. Ojha, R., A. A. Stark, H. H. Hsieh, A. P. Lane, R. A. Chamberlin, T. M. Bania, A. D. Bolatto, J. M. Jackson, and G. A. Wright. 2001. AST/ RO Observations of Atomic Carbon near the Galactic Center. Astrophysical Journal, 548: 253? 257. Oka, T., T. Hasegawa, F. Sato, M. Tsuboi, and A. Miyazaki. 1998. A Large-Scale CO Survey of the Galactic Center. Astrophysical Jour- nal, Supplement Series, 118: 455? 515. Robinson, B. J., F. F. Gardner, K. J. van Damme, and J. G. Bolton. 1964. An Intense Concentration of OH Near the Galactic Centre. Nature, 202: 989? 991. Stark, A. A., J. Bally, G. R. Knapp, and R. W. Wilson. 1988. ?The Bell Laboratories CO Survey.? In Molecular Clouds in the Milky Way and External Galaxies, ed. R. L. Dickman, R. L. Snell, and J. S. Young, p. 303. New York: Springer-Verlag. Stark, A. A., J. Bally, R. W. Wilson, and M. W. Pound. 1989. ?Molecular Line Observations of the Galactic Center Region.? In The Center of the Galaxy: Proceedings of the 136th Symposium of the Inter- national Astronomical Union, ed. M. Morris, p. 129. Dordrecht: Kluwer Academic Publishers. Stark, A. A., J. Bally, O. E. Gerhard, and J. Binney. 1991. On the Fate of Galactic Centre Molecular Clouds. Monthly Notices of the Royal Astronomical Society, 248: 14P? 17PP. Stark, A. A., J. Bally, S. P. Balm, T. M. Bania, A. D. Bolatto, R. A. Cham- berlin, G. Engargiola, M. Huang, J. G. Ingalls, K. Jacobs, J. M. Jackson, J. W. Kooi, A. P. Lane, K.-Y. Lo, R. D. Marks, C. L. Mar- tin, D. Mumma, R. Ojha, R. Schieder, J. Staguhn, J. Stutzki, C. K. Walker, R. W. Wilson, G. A. Wright, X. Zhang, P. Zimmermann, and R. Zimmermann. 2001. The Antarctic Submillimeter Telescope and Remote Observatory (AST/RO). Publications of the Astronom- ical Society of the Pacifi c, 113: 567? 585. Stark, A. A., C. L. Martin, W. M. Walsh, K. Xiao, A. P. Lane, and C. K. Walker. 2004. Gas Density, Stability, and Starbursts near the Inner Lindblad Resonance of the Milky Way. Astrophysical Journal Let- ters, 614: L41? L44. 28_Martin_pg369-372_Poles.indd 37228_Martin_pg369-372_Poles.indd 372 11/17/08 9:41:03 AM11/17/08 9:41:03 AM ABSTRACT. The High Elevation Antarctic Terahertz Telescope (HEAT) is a proposed 0.5-m THz observatory for automated, remote operation at the summit of Dome A, the highest point on the Antarctic plateau. The altitude of Dome A combined with the extreme cold and dry conditions prevalent there make it the best location on Earth for conducting many types of astronomical observations. The HEAT will operate at wave- lengths from 150 to 400 micrometers and will observe the brightest and most diagnostic spectral lines from the galaxy. It will follow PreHEAT, an NSF-funded 450-micrometer tipper and spectrometer that was deployed to Dome A in January 2008 by the Polar Research Institute of China. PreHEAT is one of several instruments designed to oper- ate with the University of New South Wales? Plateau Observatory (PLATO). A 1.5-THz (200-micrometer) receiver channel will be installed onto PreHEAT in Austral summer 2008? 2009. PreHEAT/HEAT and PLATO operate autonomously from Dome A for up to a year at a time, with commands and data being transferred to and from the experi- ment via satellite daily. The Plateau Observatory is the Dome A component of the multi- national Astronomy at the Poles (AstroPoles) program, which has been endorsed by the Joint Committee for the International Polar Year (IPY). INTRODUCTION From the Milky Way to high-redshift protogalaxies, the internal evolu- tion of galaxies is determined to a large extent by the life cycles of interstellar clouds, as shown in Figure 1. These clouds are largely comprised of atomic and molecular hydrogen and atomic helium, which are notoriously diffi cult to detect under normal interstellar conditions. Atomic hydrogen is detectable via the 21-cm spin-fl ip transition and provides the observational basis for current models of a multiphase galactic interstellar medium (ISM). Its emission is in- sensitive to gas density and does not always discriminate between cold (T H1101170 K) atomic clouds and the warm (T H110118000 K), neutral medium that is thought to pervade the galaxy. Furthermore, neither atomic helium nor molecular hy- drogen (H 2 ) have accessible emission line spectra in the prevailing physical conditions in cold interstellar clouds. Thus, it is important to probe the nature of the ISM via rarer trace elements. Carbon, for example, is found in ionized form (C H11001 ) in neutral clouds, eventually becoming atomic (C), then molecular as carbon monoxide (CO) in dark molecular clouds. Christopher Walker and Craig Kulesa, Uni- versity of Arizona, 933 North Cherry Avenue, Tucson, AZ 85721, USA. Corresponding au- thor: C. Walker (cwalker@as.arizona.edu). Ac- cepted 25 June 2008. HEAT: The High Elevation Antarctic Terahertz Telescope Christopher K. Walker and Craig A. Kulesa 29_Walker_pg373-380_Poles.indd 37329_Walker_pg373-380_Poles.indd 373 11/17/08 9:49:02 AM11/17/08 9:49:02 AM 374 SMITHSONIAN AT THE POLES / WALKER AND KULESA Although we are now beginning to understand star formation, the formation, evolution, and destruction of molecular clouds remains shrouded in uncertainty. The need to understand the evolution of interstellar clouds in the context of star formation has become a central theme of contemporary astrophysics. Indeed, the National Re- search Council?s most recent decadal survey has identifi ed the study of star formation as one of the key recommenda- tions for new initiatives in this decade. HEAT SCIENCE GOALS Via resolved C H11001 , C, CO, and N H11001 THz line emission, the High Elevation Antarctic Terahertz Telescope (HEAT) uniquely probes the pivotal formative and disruptive stages in the life cycles of interstellar clouds and sheds crucial light on the formation of stars by providing new insight into the relationship between interstellar clouds and the stars that form in them, a central component of galactic evolution. A detailed study of the ISM of the Milky Way is used to construct a template to interpret global star forma- tion in other spiral galaxies. The minimum science mission of HEAT is to make sig- nifi cant contributions to achieving the three major science goals described below. Using the proposed instrument and observing methodology, the minimum mission is expected to be achievable in a single season of survey operation from Dome A. GOAL 1: OBSERVING THE LIFE CYCLE OF INTERSTELLAR CLOUDS The formation of interstellar clouds is a prerequisite for star formation, yet the process has not yet been observed! The HEAT is designed with the unique combination of sen- sitivity and resolution needed to observe atomic clouds in the process of becoming giant molecular clouds (GMCs) and their subsequent dissolution into diffuse gas via stellar feedback. GOAL 2: MEASURING THE GALACTIC STAR FORMATION RATE The HEAT will probe the relation between the gas surface density on kiloparsec scales and the N H11001 H11002derived FIGURE 1. The High Elevation Antarctic Terahertz Telescope (HEAT) will observe the fi ne-structure lines of N H11001 , C H11001 , C, and CO that probe the entire life cycle of interstellar clouds. In particular, HEAT will witness the transformation of neutral atomic clouds into star-forming clouds, the interaction of the interstellar medium (ISM) with the young stars that are born from it, and the return of enriched stellar material to the ISM by stellar death. FIGURE 2. The location of GMCs in the nearby spiral galaxy M33 are overlaid upon an integrated intensity map of the HI 21-cm line (Engargiola et al., 2003). These observations show that GMCs are formed from large structure of atomic gas, foreshadowing the detailed study of GMC formation that HEAT will provide in the Milky Way. 29_Walker_pg373-380_Poles.indd 37429_Walker_pg373-380_Poles.indd 374 11/17/08 9:49:03 AM11/17/08 9:49:03 AM THE HEAT TELESCOPE 375 star formation rate, so that we might be able to better understand the empirical Schmidt law used to estimate the star-forming properties of external galaxies (Schmidt, 1959; Kennicutt et al., 1998). GOAL 3: CONSTRUCTING A MILKY WAY TEMPLATE C H11001 and N H11001 will be the premier diagnostic tools for terahertz studies of external galaxies with large redshifts (e.g., with Atacama Large Millimeter Array, or ALMA). In such spatially unresolved galaxies, however, only global properties can be measured. The HEAT observations will yield detailed interstellar studies of the widely varying conditions in our own Milky Way galaxy and serve as a crucial diagnostic template or ?Rosetta Stone? that can be used to translate the global properties of more distant galaxies. PROPERTIES OF THE PROPOSED SURVEY The HEAT?s science drivers represent a defi nitive sur- vey that would not only provide the clearest view of inter- stellar clouds and their evolution in the galaxy but would also serve as the reference map for contemporary focused studies with space, suborbital, and ground observatories. The following properties defi ne the science requirements for the HEAT survey. HIGH-RESOLUTION SPECTROSCOPIC IMAGING Techniques commonly used to diagnose the molecu- lar ISM include submillimeter continuum mapping of dust emission (Hildebrand et al., 1983) and dust extinction mapping at optical and near-infrared wavelengths (Lada et al., 1994). Large-format detector arrays in the infrared are now commonplace, and with the advent of bolometer arrays, both techniques have performed degree-scale maps of molecular material. However, these techniques have limited applicability to the study of the structure of the galactic ISM due to the complete lack of kinematic infor- mation. The confl uence of many clouds along most galactic lines of sight can only be disentangled with spectral line techniques. Fitting to a model of galactic rotation is often the only way to determine each cloud?s distance and loca- tion within the galaxy. With resolution fi ner than 1 km/s, a cloud?s kinematic location can even be distinguished from other phenomena that alter the line shape, such as tur- bulence, rotation, and local effects, such as protostellar outfl ows. These kinematic components play a vital role in the sculpting of interstellar clouds, and a survey that has the goal of understanding their evolution must be able to measure them. The HEAT will easily resolve the intrinsic profi les of galactic interstellar lines, with a resolution of H110210.4 km/s up to 370 km/s of spectrometer bandwidth, comparable to the galactic rotational velocity. A TERAHERTZ GALACTIC PLANE SURVEY Molecular line surveys have been performed over the entire sky in the light of the 2.6 mm J H11005 1? 0 line of 12 CO, and they have been used to synthesize our best un- derstanding of the molecular content of the galaxy. Still, our understanding of the evolution of galactic molecular clouds is woefully incomplete. As already described in the HEAT Science Goals section, the dominant spectral lines of the galaxy are the fi ne-structure far-infrared and sub- millimeter lines of C, CO, C H11001 , and N H11001 . They probe and regulate all aspects of the formation and destruction of star-forming clouds. They will provide the fi rst barometric maps of the galaxy and illuminate the properties of clouds and their life cycles in relation to their location in the gal- axy. They will highlight the delicate interplay between (massive) stars and the clouds which form them, a criti- cal component of galactic evolution. A terahertz survey will dramatically enhance the value of existing millimeter- wave CO observations by providing critical excitation constraints. ARCMINUTE ANGULAR RESOLUTION AND FULLY SAMPLED MAPS Good angular resolution is a critical aspect of im- provement for a new galactic survey. Previous surveys of [N II] and [C II] were limited to very small regions (KAO and ISO) or had low angular resolution (COBE and BICE) (Bennett et al., 1994; Nakagawa et al., 1998). The HEAT will fully sample both species over large regions of sky to their diffraction-limited resolution of 1.7H11032 and 1.3H11032, re- spectively. Arcminute resolution with proper sampling is crucial to disentangling different clouds and cloud com- ponents over large distances in the galaxy. For example, the Jeans length for star formation in a GMC is approxi- mately 0.5 pc. This length scale is resolved by HEAT to a distance of 500 pc at CO J H11005 7? 6 and [C I] and 1200 pc at [C II]. Warm and cold HI clouds and GMCs can be resolved well past 10 kpc. 29_Walker_pg373-380_Poles.indd 37529_Walker_pg373-380_Poles.indd 375 11/17/08 9:49:24 AM11/17/08 9:49:24 AM 376 SMITHSONIAN AT THE POLES / WALKER AND KULESA HIGH SENSITIVITY The HEAT?s high sensitivity is due mostly to the su- perlative atmospheric conditions expected above Dome A, Antarctica. The extreme cold and exceptional dryness al- low ground-based observations into the otherwise forbid- den terahertz windows. A plot of the expected atmospheric transmission for excellent winter observing conditions at Dome A versus the comparable opacity at the South Pole is plotted in Figure 3. The high elevation, cold atmosphere, and benign wind conditions at Dome A defi nitively open the terahertz windows to ground-based observatories and can- not be matched anywhere else on Earth. The implications for the sensitivity to each spectral line is discussed below. CO J H11005 7? 6 and [C I] J H11005 2? 1 We aim to detect all CO and C 0 to A V H11005 1? 2, where most hydrogen has formed H 2 and CO is just forming. This extinction limit corresponds to N(CO) H110115 H11003 10 15 cm H110022 and N(C) H11005 1.6 H11003 10 16 cm H110022 for integrated intensi- ties of 3 K k/ms in CO J H11005 7? 6 and 1.8 K km/s in [C I]. These sensitivity limits are achievable (three sigma) within 1.6 and 5 minutes, respectively, of integration time at 810 GHz in median winter atmospheric conditions on Dome A with an uncooled Schottky receiver. Limits on line emis- sion in that time would constrain the gas density, based upon the line brightness of millimeter wave transitions. N H11001 and C H11001 The fi ne-structure lines of ionized carbon and nitrogen represent the dominant coolants of the interstellar medium of the galaxy and star-forming galaxies. Indeed, the inte- grated intensity of the 158-micrometer C H11001 line alone rep- resents H110111% of the bolometric luminosity of the galaxy! As such, these lines are relatively easy to detect in the ISM. Our most demanding requirements for detection of C H11001 and N H11001 lie in the search for the formation of giant molecular clouds (via C H11001 ) and the measurement of the diffuse warm ionized medium in the galaxy (via N H11001 ). A fl ux limit of 2 K km/s will detect N H11001 in warm HI as far away as the molecu- lar ring, achievable in good winter weather in three minutes with velocity smoothing to 3 km/s, appropriate for hot ion- ized gas. Similarly, the accumulation of GMCs from many cold neutral clouds of atomic hydrogen occurs at low rela- tive column densities of H110115 H11003 10 20 cm H110022 . Since essentially all carbon in such clouds is ionized, N(C H11001 ) H1101110 17 cm H110022 . At the T H11005 70 K common in cold atomic clouds and n H H11005 10 3 cm H110023 , the expected C H11001 line emission would be 2.5 K km/s, detectable with a Schottky receiver in 10 minutes in excellent winter weather on Dome A. The three sigma limit achievable with deep integrations (two hours) with HEAT would reach n H H11005 10 2 cm H110023 . This pressure limit would readily determine whether interstellar material causing sig- nifi cant infrared extinction but without CO is gravitation- ally bound and likely to be a forming molecular cloud or is simply a line of sight with numerous overlapping diffuse HI clouds. LARGE-AREA MAPPING COVERAGE OF THE GALACTIC PLANE From previous CO surveys it is known that the scale height of CO emission toward the inner galaxy is less than one degree (Dame et al., 1987, 2001). The BICE balloon experiment demonstrated that the C H11001 distribution is more extended but is still confi ned to ?b? H11021 1. Interstellar pres- sure, abundances, and physical conditions vary strongly as a function of galactocentric radius, so it is necessary to probe the inner galaxy, the outer galaxy and both spiral arms and interarm regions to obtain a statistically mean- ingful survey that encompasses the broad dynamic range of physical conditions in the galaxy. We propose therefore to probe the entire galactic plane as seen from Dome A (0 o H11022 l H11022 H11002120 o ). An unbiased survey will be undertaken, ul- FIGURE 3. Each of HEAT?s heterodyne beams is overlaid upon a 2MASS infrared image of NGC 6334. The beams will measure high- resolution spectra in the 0.81-, 1.46-, and 1.90-THz bands, respec- tively; a small portion (25%) of each is shown as synthetic spectra of NGC 6334. 29_Walker_pg373-380_Poles.indd 37629_Walker_pg373-380_Poles.indd 376 11/17/08 9:49:25 AM11/17/08 9:49:25 AM THE HEAT TELESCOPE 377 timately covering up to 240 square degrees (H110021 o H11021 b H11021 1 o ); however, 80 square degrees in two years will be targeted by the Schottky receiver system described here. Figure 4 dem- onstrates the sky coverage of HEAT?s survey of the inner galaxy, with the fi rst season coverage highlighted in yel- low. It will probe three crucial components of the galaxy: the molecular ring, the Crux spiral arm, and the interarm region. The remaining sky coverage will be provided by a future upgraded instrument package from the Netherlands Institute for Space Research (SRON), featuring a cryo- cooled 4 K superconductor insulator superconductor and hot-electron bolometer system. The ?inner? galaxy survey will coincide with Galactic Legacy Infrared Mid-Plane Sur- vey Extraordinaire (GLIMPSE), a Spitzer Space Telescope (SST) Legacy Program (Benjamin et al., 2003). Above l H11005 90 o , most of the CO emission is located at higher galactic latitude, so l and b ?strip mapping? will locate the target regions, generally following the outskirts of CO J H11005 1? 0 distribution (Dame et al., 1987, 2001), and the best-char- acterized star-forming regions in the galaxy. The observing program will be designed to maximize synergies with the ?Cores to Disks? SST Legacy program (Evans et al., 2003) and other SST GTO programs. HEAT INSTRUMENTATION OVERVIEW The HEAT will be a fully automated, state-of-the-art terahertz observatory designed to operate autonomously from Dome A in Antarctica. The combination of high alti- tude (4,100 m), low precipitation, and extreme cold make the far-infrared atmospheric transmission exceptionally good from this site. In Figure 5 we present a plot of the ex- pected atmospheric transmission above Dome A as a func- tion of wavelength (Lawrence, 2004), indicating that win- ter weather at Dome A approaches (to order of magnitude) the quality of that achieved by the Stratospheric Observa- tory for Infrared Astronomy (SOFIA). The wavelengths of several important astrophysical lines are indicated with arrows. The HEAT is designed to take advantage of these unique atmospheric conditions and observe simultane- ously in [C II] (158 micrometer), [N II] (205 micro meter), and CO J H11005 7? 6 and [C I] (370 micrometer). A conceptual drawing of HEAT is shown in Figure 6. For robustness and effi ciency, the telescope and instrument are integrated into a common optical support structure. The HEAT will be mounted on top of the University of New South Wales? Plateau Observatory (PLATO), which was deployed to Dome A in January 2008. The Plateau Observatory is the successor to the Automated Astrophys- ical Site-Testing International Observatory (AASTINO) deployed to Dome C in 2003, and it provides power and FIGURE 4. An 8.3-micrometer map of the galactic plane from the molecular ring through the Scutum-Crux spiral arm (-20 o H11022 l H11022 -55 o ). The yellow rectangle highlights the region to be explored by HEAT in its fi rst season at Dome A. A defi nitive chemical and kinematic survey of star- forming clouds in [C I] J H11005 2? 1 and 12 CO J H11005 7? 6 of 40 square degrees (H1101110 square degrees in [C II] and [N II] emission) can be performed in a single season using Schottky receivers. No other site on Earth allows routine access to both far-infrared lines. FIGURE 5. Terahertz atmospheric transmission for good (twenty- fi fth percentile) winter conditions for the South Pole (bottom line) and Dome A (top line), derived from PWV measurements at the South Pole, atmospheric models from Lawrence (2004), and actual automatic weather station data collected during 2005 from Dome A. The PWV content for each model atmosphere is 210 and 50 mi- crometers, respectively. Arrows indicate the wavelengths of the [N II], [C II], and CO/[C I] lines. 29_Walker_pg373-380_Poles.indd 37729_Walker_pg373-380_Poles.indd 377 11/17/08 9:49:39 AM11/17/08 9:49:39 AM 378 SMITHSONIAN AT THE POLES / WALKER AND KULESA communications for the HEAT telescope and instrument. The total power budget for HEAT, including cryogenics, telescope drive system, and instrument control system, is maximally 600 W, which is readily provided by effi cient, high-reliability generators within PLATO. Data transfer and control of HEAT will be done via Iridium satellite through the PLATO facilities. The combined HEAT and PLATO facility is functionally equivalent to a space-based observatory. The HEAT will be able to calibrate observations through several means. (1) A vane with an ambient tem- perature absorbing load will be located at the cryostat en- trance window, allowing standard chopper wheel calibra- tion to be performed. (2) HEAT will routinely perform sky dips to compute the atmospheric optical depth in each of its three wavelength bands. (3) HEAT will regularly ob- serve a standard list of calibration sources. (4) The PLATO currently hosts PreHEAT, a 450-micrometer tipper and spectrometer that measures atmospheric transmission. Its measurements will be coordinated with HEAT spectral line observations to provide cross calibration. LOGISTICS: DEPLOYMENT TO DOME A Antarctic science has reached a level of maturity where several options exist for fi elding instruments on remote sites. For Dome A, these options include the following: 1. (Chinese) Traverse from Zhongshan Station to Dome A: PLATO and its complement of instruments (includ- ing PreHEAT) were deployed to Dome A by a 1300-km Chinese traverse from the coastal Zhongshan station in January 2008. The expedition was a collaborative effort with the Polar Research Institute of China, the National Astronomical Observatory of China, and the Nanjing Institute of Astronomical Optics Technology. Upgrades to PreHEAT and the installation of the full HEAT experiment could potentially be deployed in a similar manner. 2. (American) Twin Otter air support: If a Chinese tra- verse brings PLATO (in 2007? 2008) and HEAT to Dome A (in 2009? 2010), U.S. Antarctic Program Twin Otter air support would allow personnel to be fl own in from the South Pole or the forthcoming AGAP fi eld camps (such as AGO3) to facilitate the HEAT instal- lation. Furthermore, the HEAT experiment is small enough to be deployed by Twin Otter. 3. (Australian) An Australian Antarctic Division CASA 212 cargo fl ight directly from Mawson/Davis to Dome A may be possible for transport of the HEAT experi- ment, with subsequent fl ights to support fuel and/or personnel for the HEAT installation. SUMMARY At this writing, the pathfi nder for HEAT, PreHEAT, is operating autonomously from a PLATO module recently deployed to the summit of Dome A by the Polar Research Institute of China. Because of the altitude and extreme cold/dry conditions known to exist at Dome A, the atmo- FIGURE 6. The HEAT telescope has an effective collecting area of 0.5 m. Elevation tracking is accomplished by rotating the 45? fl at refl ector. The entire telescope structure is warmed by waste heat from the PLATO instrument module below. The Schottky mixers used in the instrument package are effi ciently cooled to 70 K using a reliable off-the-shelf closed-cycle cryocooler. 29_Walker_pg373-380_Poles.indd 37829_Walker_pg373-380_Poles.indd 378 11/17/08 9:49:51 AM11/17/08 9:49:51 AM THE HEAT TELESCOPE 379 spheric opacity above the site is expected to be the lowest on Earth, making it ideal for far-infrared/terahertz obser- vatories. Our hope is that HEAT will follow quickly on the heels of PreHEAT and provide a powerful new win- dow to the universe. LITERATURE CITED Bennett, C. L., D. J. Fixsen, G. Hinshaw, J. C. Mather, S. H. Moseley, E. L. Wright, R. E. Eplee Jr., J. Gales, T. Hewagama, R. B. Isaacman, R. A. Shafer, and K. Turpie. 1994. Morphology of the Interstellar Cooling Lines Detected by COBE. Astrophysical Journal, 434: 587? 598. Benjamin, R. A., E. Churchwell, B. L. Babler, T. M. Bania, D. P. Clemens, M. Cohen, J. M. Dickey, R. Indebetouw, J. M. Jackson, H. A. Kobulnicky, A. Lazarian, A. P. Marsten, J. S. Mathis, M. R. Meade, S. Seager, S. R. Stolovy, C. Watson, B. A. Whitney, M. J. Wolff, and M. G. Wolfi re. 2003. GLIMPSE: I. An SIRTF Legacy Project to Map the Inner Galaxy. Publications of the Astronomical Society of the Pacifi c, 115: 953? 964. Dame, T. M., H. Ungerechts, R. S. Cohen, E. J. Geus, de, I. A. Grenier, J. May, D. C. Murphy, L.-? Nyman, and P. Thaddeus. 1987. A Com- posite CO Survey of the Entire Milky Way. Astrophysical Journal, 322: 706. Dame, T. M., D. Hartmann, and P. Thaddeus. 2001. The Milky Way in Molecular Clouds: A New Complete CO Survey. Astrophysical Journal, 547: 792. Engargiola, G., R. L. Plambeck, E. Rosolowsky, and L. Blitz. 2003. Giant Molecular Clouds in M33. I. BIMA All-Disk Survey. Astrophysical Journal, Supplement Series, 149: 343? 363. Evans, N. J., II, and the Cores to Disks (c2d) Team. 2003. From Molecu- lar Cores to Planet-forming Disks: A SIRTF Legacy Program. Publi- cations of the Astronomical Society of the Pacifi c, 115: 965? 980. Hildebrand, R. H. 1983. The Determination of Cloud Masses and Dust Characteristics from Submillimetre Thermal Emission. Quarterly Journal of the Royal Astronomical Society, 24: 267. Kennicutt, R. C. 1998. The Global Schmidt Law in Star-Forming Galax- ies. Astrophysical Journal, 498: 541. Lada, C. J., E. A. Lada, D. P. Clemens, and J. Bally. 1994. Dust Extinc- tion and Molecular Gas in the Dark Cloud IC 5146. Astrophysical Journal, 429: 694. Lawrence, J. S. 2004. Infrared and Submillimetre Atmospheric Charac- teristics of High Antarctic Plateau Sites. Publications of the Astro- nomical Society of the Pacifi c, 116: 482. Nakagawa, T., Y. Y. Yui, Y. Doi, H. Okuda, H. Shibai, K. Mochizuki, T. Nishimura, and F. J. Low. 1998. Far-Infrared [Cii] Line Survey Observations of the Galactic Plane. Astrophysical Journal, Supple- ment Series, 115: 259? 269. Schmidt, M. 1959. The Rate of Star Formation. Astrophysical Journal, 129: 243. 29_Walker_pg373-380_Poles.indd 37929_Walker_pg373-380_Poles.indd 379 11/17/08 9:50:00 AM11/17/08 9:50:00 AM 29_Walker_pg373-380_Poles.indd 38029_Walker_pg373-380_Poles.indd 380 11/17/08 9:50:00 AM11/17/08 9:50:00 AM ABSTRACT. Astronomical instruments on the Antarctic plateau are very well suited to observing the formation of stars and their associated planetary systems since young stars emit their light at the wavelengths at which Antarctica offers the most striking ad- vantages. Antarctic telescopes have already brought new insights into the physics of star formation and the molecular clouds where it occurs. During the International Polar Year (IPY), new sites will be opened up to astronomical exploitation, with the prospect of new capabilities in the drive to understand how stars and planets form. INTRODUCTION Stars are one of the main engines of evolution in the universe. They convert mass to light and hydrogen and helium into heavier elements; massive stars compress and disrupt nearby gas clouds by the action of their ionizing radiation and their stellar winds. However, the formation and early evolution of stars are not well understood: they form inside clouds of molecular gas and dust, which are opaque to visible light but transparent to infrared light and submillimeter- wave radiation. These wavebands are thus crucial to our understanding of the formation of stars: the young stars themselves radiate infrared light, which can penetrate the dark clouds, while submillimeter-wave observations can trace the gas and dust that make up the clouds. ANTARCTIC SUBMILLIMETER TELESCOPE AND REMOTE OBSERVATORY OBSERVATIONS OF MOLECULAR CLOUDS The main constituents of molecular clouds, hydrogen and helium gases, are effectively invisible to us: both molecular hydrogen and atomic helium have very few low-energy transitions that could be excited at the low temperatures prevailing in interstellar space. We therefore rely on tracers? gas and dust that are readily excited at low temperatures and readily emit at long wavelengths. The most basic of these tracers is carbon monoxide (CO), which is the most abundant molecule in these gas clouds after hydrogen (H 2 ) and helium (He). N. F. H. Tothill, School of Physics, University of Exeter, Stocker Road, Exeter EX4 4QL, UK; also Smithsonian Astrophysical Observatory, 60 Garden Street, Cambridge, MA 02138, USA. M. J. McCaughrean, School of Physics, University of Exeter, Stocker Road, Exeter EX4 4QL, UK. C. K. Walker and C. Kulesa, Steward Observatory, Uni- versity of Arizona, Tucson, AZ 85721, USA. A. Loehr, Smithsonian Astrophysical Observatory, 60 Garden Street, Cambridge, MA 02138, USA. S. Parshley, Department of Astronomy, Cornell University, Ithaca, NY 14853, USA. Correspond- ing author: N. F. H. Tothill (nfht@astro.ex.ac.uk). Accepted 25 June 2008. Watching Star Birth from the Antarctic Plateau N. F. H. Tothill, M. J. McCaughrean, C. K. Walker, C. Kulesa, A. Loehr, and S. Parshley 30_Tothill_pg381-386_Poles.indd 38130_Tothill_pg381-386_Poles.indd 381 11/17/08 9:50:51 AM11/17/08 9:50:51 AM 382 SMITHSONIAN AT THE POLES / TOTHILL ET AL. The lower-energy (low-J) transitions of CO emit radiation at wavelengths of 0.8? 3mm, and are easily detected from high, dry, mountaintop sites in the temperate zones. At shorter wavelengths, the higher energy mid-J transitions can be detected only through a very dry atmosphere. The Antarctic plateau provides the largest fraction of such dry weather of any observing site in the world, and, sited at Amundsen-Scott South Pole Station, the Antarctic Sub- millimeter Telescope and Remote Observatory (AST/RO; Stark et al., 2001) is designed to take full advantage of these conditions. The mid-J transitions (from CO 4? 3 up to CO 7? 6) are particularly interesting for their ability to probe the physi- cal conditions of the molecular gas from which they arise: in particular, they can only be excited in gas with density comparable to a critical density (about 10 4 molecules cm H110023 for CO 4? 3); emission in these transitions implies the pres- ence of dense gas. By also observing the molecular clouds in lower-J transitions, AST/RO is able to trace the velocity structure of the gas clouds and to estimate the gas tem- perature, thus providing a suite of measurements of the physical conditions of the gas clouds where stars form. NEARBY LOW-MASS STAR-FORMING REGIONS All the nearest star-forming molecular clouds (within a few hundred parsecs; a parsec (pc) is a standard as- tronomical distance unit: 1 pc H11005 3.09 H11003 10 16 m H11005 3.26 light-years) form low-mass stars. Because of their proxim- ity, they subtend large areas on the sky (of the order of square degrees), requiring large amounts of time to map them properly. The Antarctic Submillimeter Telescope and Remote Observatory is very well suited to this task: its small mirror gives it a comparatively large beam, which, in turn, allows large areas to be mapped quickly. The highly transparent Antarctic atmosphere provides long stretches of very clear air in winter, which allows large blocks of time to be allocated to mapping these clouds at comparatively high frequencies. The clouds themselves are less dense and cooler than the giant molecular clouds that form the majority of stars, and many of the stars form in isolation, rather than in clusters. It is therefore often assumed that mid-J transitions of CO are not excited in these regions and are so diffi cult to detect that they no lon- ger make good tracers. The AST/RO mapped large areas of two nearby cloud complexes, Lupus and Chamaeleon (named after the constellations in which they are found), in the CO 4? 3 transition, fi nding signifi cant emission. With a brightness temperature of the order of 1K, this CO 4? 3 emission must come from molecular gas that is dense enough to thermalize the transition and warm enough to have a Rayleigh-Jeans temperature of a few Kelvin. The Lupus star-forming region consists of a complex of molecular clouds lying about 150 pc from Earth, associ- ated with a large number of young stars. The clouds are readily visible in optical photographs of the sky as clumpy, fi lamentary dark patches (Figure 1)? indeed, this is how these clouds were fi rst discovered (Barnard, 1927). The Lupus complex lies to one side of the Scorpius? Centaurus OB Association (Sco-Cen), a huge collection of very mas- sive young stars lying in the southern sky. On the other side of Sco-Cen, the rho Ophiuchi star-forming region displays clusters of massive young stars whose interac- tion with their natal molecular gas produces highly vis- ible nebulae. By comparison to rho Ophiuchi, Lupus is quiescent? it is rather lacking in massive stars and has no large cluster, the young stars being much more spread out. One might therefore assume that Lupus would also lack the dense, warm molecular gas found in abundance in rho Ophiuchi. However, the AST/RO data show detectable CO 4? 3 emission throughout the Lupus clouds, with very strong emission in a few hot spots. By comparing this emission to the more easily excited 13 CO 2? 1 (Figure 2), it is possible to estimate the physical conditions of the gas (Figure 3). The gas making up the bulk of the clouds is quite warm (prob- ably H1102210K) and close to the critical density. The clumps within the cloud seem to be denser but not much cooler, and some are warm (around 20K). The data suggest that one of the hot spots in Lupus III is very warm (perhaps as much as 50K) but not dense enough to fully excite the 4? 3 transition. The hot spot at the northwestern end of the Lu- pus I fi lament also appears to be warm and not very dense but has broader lines, implying more turbulent motion in the gas. While the elevated temperature in Lupus III can be explained by the proximity of the fairly massive young stars HR 5999 and 6000 (visible in Figure 1, lying in the dark cloud), there are no comparable stars near the end of the Lupus I fi lament. Clearly, the Lupus clouds show sig- nifi cant diversity in their physical conditions and, hence, in the environments in which stars are formed. Being more easily excited, emission in the isotopically substituted 13 CO 2? 1 line is distributed throughout the mo- lecular gas and is less discriminating as a probe of the physi- cal conditions of the gas. But because it is so widely distrib- uted, it is an excellent tracer of the velocity fi eld of the gas. The ability of AST/RO to map large areas allows us to fully sample the gas velocity fi eld over degree-scale fi elds. Maps of the centroid velocity of the 13 CO 2? 1 line show strong ve- locity gradients in several locations in the complex, usually 30_Tothill_pg381-386_Poles.indd 38230_Tothill_pg381-386_Poles.indd 382 11/17/08 9:50:52 AM11/17/08 9:50:52 AM STAR BIRTH FROM THE ANTARCTIC PLATEAU 383 FIGURE 1. Two of the molecular clouds that make up the Lupus complex, visible as dark patches against the stars: (left) Lupus I and (right) Lupus III. The ridge in Lupus I is about 5 pc long. Images taken from the Digital Sky Survey (Lasker et al., 1990). FIGURE 2. The same clouds as in Figure 1, mapped in the millimeter-wave emission of 13 CO 2? 1. The isotopically substituted CO traces the dark cloud structure very well (N. F. H. Tothill, A. Loehr, S. C. Parshley, A. A. Stark, A. P. Lane, J. I. Harnett, G. Wright, C. K. Walker, T. L. Bourke, and P. C. Myers, unpublished manuscript). 30_Tothill_pg381-386_Poles.indd 38330_Tothill_pg381-386_Poles.indd 383 11/17/08 9:50:52 AM11/17/08 9:50:52 AM 384 SMITHSONIAN AT THE POLES / TOTHILL ET AL. over quite small distances (about 0.5 pc), but in Lupus I, a different pattern emerges, of a shallow and rather uneven velocity gradient along the fi lament, coupled with a strong gradient across the fi lament. This gradient across the fi la- ment is coherent over at least 2 pc of the fi lament?s length. The presence of such a large coherent structure in a region whose activity appears quite stochastic is remarkable; it may have arisen from external infl uences, presumably from Sco-Cen. The massive stars in Sco-Cen are quite capable of affecting nearby clouds; the rho Ophiuchi region on the other side of Sco-Cen is clearly infl uenced by the nearby OB association. There are some signs of a supernova remnant to the northwest of the Lupus I fi lament, which could have a strong dynamical effect on the gas. NEW OBSERVING SITES IN THE ANTARCTIC AND ARCTIC The factors that make the South Pole such a good site to detect the submillimeter-wave radiation from the clouds around young stars are likely to be found in many other locations on the Antarctic plateau and even at some sites in the Arctic. With the astronomical potential of the Ant- arctic plateau established, it is possible to evaluate other sites in the polar regions as potential observatories. Projects to evaluate the potential of several polar sites are taking place during the International Polar Year: As- tronomy from the Poles (AstroPoles) is a general program to study all the sites listed below, while STELLA ANT- ARCTICA concentrates on a more detailed study of the characteristics of the Franco-Italian Concordia station on Dome C. The meteorology of Antarctica is dominated by the katabatic fl ow, as the air loses heat by contact with the radiatively cooled snow surface, loses buoyancy, and sinks down the slope of the ice sheet, leading to a downward- fl owing wind. Most of the atmospheric turbulence is con- centrated in a boundary layer between this fl ow and the ice, a layer which gets deeper farther downslope. ANTARCTICA: DOME A Dome A is the highest point on the plateau and has less atmosphere to get in the way than any other site. How- ever, it also lacks infrastructure: in the absence of a year- round station, all instruments must be fully autonomous. Traverses to Dome A and astronomical site testing are be- ing undertaken by the Polar Research Institute of China, in collaboration with an international consortium. This consortium, including Chinese institutions, the Universi- ties of New South Wales, Arizona, and Exeter, and Caltech, FIGURE 3. Comparison of CO 4? 3 and 13 CO 2? 1 emission from Lupus I and III clouds. The different physical conditions of the hot spots in the two clouds show up in the different distributions, but the bulk of each cloud is quite similar to that of the other (N. F. H. Tothill, A. Loehr, S. C. Parshley, A. A. Stark, A. P. Lane, J. I. Harnett, G. Wright, C. K. Walker, T. L. Bourke, and P. C. Myers, unpublished manuscript). 30_Tothill_pg381-386_Poles.indd 38430_Tothill_pg381-386_Poles.indd 384 11/17/08 9:51:20 AM11/17/08 9:51:20 AM STAR BIRTH FROM THE ANTARCTIC PLATEAU 385 is constructing and deploying the Plateau Observatory (PLATO), an automated unmanned site-testing observatory (Lawrence et al., 2006). One of the instruments on PLATO, PreHEAT, is designed to characterize the site quality at sub- millimeter wavelengths and to map the J H11005 6? 5 emission of isotopically substituted 13 CO from massive star-forming regions and giant molecular clouds. It will be succeeded by the High Elevation Antarctic Terahertz Telescope (HEAT), a 0.5-m-aperture telescope designed to function around 0.2 mm wavelength. The HEAT will map the fi ne-structure emission from atomic and ionized carbon and nitrogen, to- gether with CO 7? 6, to trace the evolution and recycling of the interstellar medium in our galaxy. The prospects for PreHEAT and HEAT at Dome A are discussed in more de- tail in Walker and Kulesa (2009, this volume). At the top of the continent, the boundary layer at Dome A may be only 3 to 4 m thick, which would make it much easier to place a telescope above the boundary layer, into what is likely to be very stable air with very good see- ing. Other experiments on PLATO will test these predic- tions about the boundary layer at Dome A and sketch out its potential as a future observatory site. ANTARCTICA: DOME C Dome C is signifi cantly higher than the South Pole (around 3200 m elevation) and lies on a local maximum of the ice sheet along the ridge running through East Ant- arctica. It is likely to enjoy better conditions than the South Pole but not such good conditions as Dome A. Recent measurements (Lawrence et al., 2004) show the boundary layer to be about 30 m deep, with clear-air seeing above the layer estimated to be better than 0.3 arcseconds. Although Dome C probably enjoys better submillimeter atmospheric transparency than the South Pole, infrared astronomy seems more likely to be its greatest strength: the combina- tion of excellent seeing, very little cloud cover, low ther- mal background (due to the cold air), and the presence of a year-round crewed station with strong logistical support makes it possible to build an infrared facility telescope with a primary mirror diameter of 2 m or more, which would be competitive with the largest telescopes elsewhere in the world (Burton et al., 2005). Detailed site testing at Con- cordia is being carried out by the IPY program STELLA ANTARCTICA and by members of the European Commis- sion? funded network ?Antarctic Research?A European Network for Astronomy? (ARENA). In the near future, this should lead to the construction and validation of a model of the boundary layer that allows telescopes to be designed to take advantage of the unusual conditions at this site. Two small telescopes are in the process of being designed and deployed to Concordia: Antarctica Search for Transiting Extrasolar Planets (ASTEP) is an optical time series experi- ment, designed to monitor the brightness of nearby stars and watch for the fl uctuations as their planets transit. The International Robotic Antarctic Infrared Telescope (IRAIT) is a 0.8-m-aperture infrared telescope to carry out wide- fi eld infrared surveys and to test the site characteristics for a larger infrared telescope. GREENLAND: SUMMIT The atmosphere above the Greenland ice cap in winter is also dominated by katabatic fl ow. The peak of the ice cap (Summit station, at latitude 72?N) is therefore analogous to Dome A in Antarctica, albeit less extreme: lower (3200 m), warmer (about H1100242?C in good observing conditions), etc. Nonetheless, it should share many characteristics of Antarctic plateau sites, such as cold, dry, stable air. It has its own advantages: it is crewed year-round and access is easy in summer, possible in winter; it also has access to the northern sky, which is invisible from Antarctica. All these factors combine to give Summit excellent potential as an observatory site. There is also room for synergy with Antarctic observatories in order to cover more of the sky and to prototype and test equipment at a more easily ac- cessible location. ELLESMERE ISLAND The northern tip of Ellesmere Island in Canada lies very close to the North Pole and thus offers at least one of the advantages of very high latitude sites, namely, the ready availability of 24-hour darkness with small changes in source elevation. Since this part of Ellesmere Island is rocky, with mountain peaks rising above the permanent sea ice, it is likely to have rather different meteorology to the smooth ice caps in Greenland and Antarctica. Auto- mated weather stations have been placed on several can- didate rocky peaks on the northern coast, and the fi rst re- sults of this basic site testing are expected shortly. CONCLUSIONS Understanding the process of star formation requires the observation of infrared light (to see the young star through the gas and dust around it) and submillimeter waves (to estimate the physical conditions of the molec- ular gas itself). Observations from the Antarctic plateau 30_Tothill_pg381-386_Poles.indd 38530_Tothill_pg381-386_Poles.indd 385 11/17/08 9:51:23 AM11/17/08 9:51:23 AM 386 SMITHSONIAN AT THE POLES / TOTHILL ET AL. offer large advantages in both of these regimes, demon- strated by the AST/RO observations of nearby molecular clouds, which yield a picture of quite different molecular clouds: Lupus I has strong coherent velocity gradients and may have been externally infl uenced, while Lupus III has been heated by associated young stars. During the International Polar Year, efforts to test the suitability of other sites for astronomy are under way: Concordia Station on Dome C is likely to be an excellent infrared site and is the best known. Other sites (Dome A, Summit, and Ellesmere Island) are at much earlier stages of characterization. ACKNOWLEDGMENTS Arctic astronomical projects are being coordinated, for Ellesmere Island by Ray Carlberg (University of Toronto) and Eric Steinbring (Herzberg Institute of Astrophysics), and for Greenland by Michael Andersen (Niels Bohr In- stitute). The AstroPoles and STELLA ANTARCTICA pro- grams are led by Michael Burton (University of New South Wales) and Eric Fossat (University of Nice), respectively. The ARENA network is coordinated by Nicolas Epchtein (University of Nice) and funded by the European Commission under the 6th Framework Programme. Op- erations at Dome A are carried out by the Polar Research Institute of China. The AST/RO was supported by the National Science Foundation?s Offi ce of Polar Programs under ANT-0441756. We also acknowledge fi nancial sup- port from the University of Exeter. LITERATURE CITED Barnard, E. E. 1927. Catalog of 327 Dark Objects in the Sky. Chicago: University of Chicago Press. Burton, M. G., J. S. Lawrence, M. C. B. Ashley, J. A. Bailey, C. Blake, T. R. Bedding, J. Bland-Hawthorn, I. A. Bond, K. Glazebrook, M. G. Hidas, G. Lewis, S. N. Longmore, S. T. Maddison, S. Mattila, V. Minier, S. D. Ryder, R. Sharp, C. H. Smith, J. W. V. Storey, C. G. Tinney, P. Tuthill, A. J. Walsh, W. Walsh, M. Whiting, T. Wong, D. Woods, and P. C. M. Yock. 2005. Science Programs for a 2-m Class Telescope at Dome C, Antarctica: PILOT, the Pathfi nder for an International Large Optical Telescope. Publications of the Astro- nomical Society of Australia, 22: 199? 235. Lasker, B. M., C. R. Sturch, B. J. McLean, J. L. Russell, H. Jenker, and M. M. Shara. 1990. The Guide Star Catalog. I? Astronomi- cal Foundations and Image Processing. Astronomical Journal, 99: 2019? 2058, 2173? 2178. Lawrence, J. S., M. C. B. Ashley, A. Tokovinin, and T. Travouillon. 2004. Exceptional Astronomical Seeing Conditions above Dome C in Antarctica. Nature, 431: 278? 281. Lawrence, J. S., M. C. B. Ashley, M. G. Burton, X. Cui, J. R. Everett, B. T. Indermuehle, S. L. Kenyon, D. Luong-Van, A. M. Moore, J. W. V. Storey, A. Tokovinin, T. Travouillon, C. Pennypacker, L. Wang, and D. York. 2006. Site Testing Dome A, Antarctica. Proceedings of SPIE, 6267: 51? 59. Stark, A. A., J. Bally, S. P. Balm, T. M. Bania, A. D. Bolatto, R. A. Chamberlin, G. Engargiola, M. Huang, J. G. Ingalls, K. Jacobs, J. M. Jackson, J. W. Kooi, A. P. Lane, K.-Y. Lo, R. D. Marks, C. L. Martin, D. Mumma, R. Ojha, R. Schieder, J. Staguhn, J. Stutzki, C. K. Walker, R. W. Wilson, G. A. Wright, X. Zhang, P. Zimmer- mann, and R. Zimmermann. 2001. The Antarctic Submillimeter Telescope and Remote Observatory (AST/RO). Publications of the Astronomical Society of the Pacifi c, 113: 567? 585. Walker, C. K., and C. A. Kulesa. 2009. ?HEAT: The High Elevation Ant- arctic Terahertz Telescope.? In Smithsonian at the Poles: Contribu- tions to International Polar Year Science, ed. I. Krupnik, M. A. Lang, and S. E. Miller, pp. 373?380. Washington, D.C.: Smithsonian Insti- tution Scholarly Press. 30_Tothill_pg381-386_Poles.indd 38630_Tothill_pg381-386_Poles.indd 386 11/17/08 9:51:24 AM11/17/08 9:51:24 AM ABSTRACT. The collection of meteorites from the Antarctic plateau has changed from a scientifi c curiosity to a major source of extraterrestrial material. Following initial me- teorite recoveries in 1976, the U.S. National Science Foundation, the National Aeronau- tics and Space Administration (NASA), and the Smithsonian Institution formed the U.S. Antarctic Meteorite program for the collection, curation, classifi cation, and distribution of Antarctic meteorites, which was formalized in 1981. The Smithsonian provides clas- sifi cation and serves as the long-term curatorial repository, resulting in explosive growth of the Smithsonian meteorite collection. After 30 fi eld seasons, more than 80% of the Smithsonian collection now originates from Antarctica. In addition to curation and clas- sifi cation, Smithsonian staff provide administrative leadership to the program, serve on fi eld expeditions, and provide specimens for outreach and display. Given the relatively pristine state and ancient terrestrial ages of these meteorites, they provide perhaps our best sampling of the material in our solar system. Meteorites from the Moon were fi rst recognized among the Antarctic meteorites in 1981, as was the fi rst martian meteorite the next year. In 1996, debate erupted about evidence for past microbial life in an Antarctic martian meteorite, and that debate spurred the launch of two rovers to explore Mars. Among meteorites thought to have originated on asteroids, ingredients for ancient life may have survived much higher temperatures than previously envisioned during early planetary melting and differentiation. The ongoing collection of Antarctic meteorites will enrich the scientifi c community and Smithsonian Institution in specimens and knowledge about our solar system. METEORITES FROM ANTARCTICA Serendipitous fi nds of meteorites from Antarctica were documented as early as 1912 (Adelie Land), and several such fi nds occurred in the early 1960s as sci- entifi c investigations in Antarctica increased (Lazarev, 1961; Thiel Mountains, 1962; Neptune Mountains, 1964). In 1969, with the recovery of nine meteorites in the Yamato Mountains by Japanese glaciologists, meteorites went from being mere curiosities to becoming a focus of exploration. While most accumulations of multiple meteorites represent a single fall that broke up in the atmosphere and showered an area with stones, this discovery suggested a unique concentration mechanism. These nine meteorites represented six different types, including two rare chondrites (primitive meteorites formed in the solar nebula) and a diogenite Timothy J. McCoy, Linda C. Welzenbach, and Catherine M. Corrigan, Department of Mineral Sciences, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560- 0119, USA. Corresponding author: T. J. McCoy (mccoyt@si.edu). Accepted 25 June 2008. Antarctic Meteorites: Exploring the Solar System from the Ice Timothy J. McCoy, Linda C. Welzenbach, and Catherine M. Corrigan 31_McCoy_pg387-394_Poles.indd 38731_McCoy_pg387-394_Poles.indd 387 11/17/08 9:55:58 AM11/17/08 9:55:58 AM 388 SMITHSONIAN AT THE POLES / MCCOY, WELZENBACH, AND CORRIGAN (a rock formed by melting on the surface of an asteroid) (Shima and Shima, 1973). The concentration mechanism (Figure 1) is tied to the 12 million km 2 of Antarctic ice sheet, which acts as an ideal catchment area for fallen meteorites (Harvey, 2003). As the East Antarctic ice sheet fl ows toward the margins of the continent, its progress is occasionally blocked by mountains or obstructions below the ice. In these areas, old, deep, blue ice is pushed to the surface, carrying the meteorites along with it. Strong katabatic winds cause massive defl ation, re- moving large volumes of ice and preventing the accumula- tion of snow on the stranded deposits of meteorites. The end result is a representative sampling of meteorite falls. Of additional signifi cance is the terrestrial residence time of these rocks. Antarctic meteorites record terres- trial ages ranging from tens of thousands to two million years (Welten et al., 1997) and yet are less weathered than meteorites found in temperate climates. The newly fallen meteorites are quickly frozen and preserved into the thick- ening ice sheet, reducing the amount of weathering and contamination. The relatively pristine state of the samples allows studies that were previously diffi cult or impossible. The lack of weathering also means that much smaller me- teorites survive and thus provide a broader sample of the material in our solar system. ANTARCTIC METEORITE PROGRAM The Japanese began regular collecting expeditions to the Antarctic in 1973, collecting a modest 12 mete- orites. In 1974, they returned hundreds of meteorites. During this same period, University of Pittsburgh mete- orite scientist Bill Cassidy submitted three proposals to the National Science Foundation (NSF) to fund a U.S. expedition to fi nd other suitable areas of meteorite ac- cumulation. When word of the Japanese success fi nally reached the NSF, after it had rejected the three previously submitted proposals, support was granted for a 1976? 1977 expedition. Cassidy was joined by Ed Olsen (Field Museum, Chicago) and Keizo Yanai (National Institute for Polar Research, Tokyo) to search in areas accessible by helicopter from McMurdo Station to Allan Hills. Nine specimens were found that season. These early days of Antarctic meteorite collection are wonderfully recounted in Cassidy (2003). The meteorites were curated by Olsen FIGURE 1. Diagram illustrating the mechanism by which Antarctic meteorites are concentrated in specifi c locations . See text for description of this process. 31_McCoy_pg387-394_Poles.indd 38831_McCoy_pg387-394_Poles.indd 388 11/17/08 9:55:59 AM11/17/08 9:55:59 AM U.S. ANTARCTIC METEORITE PROGRAM 389 at the Field Museum, and pieces were distributed in an ad hoc fashion to the research community. Despite the modest numbers for this joint U.S.? Japanese team, it was clear that this was merely the tip of the iceberg and that large numbers of meteorites from the cleanest environment on Earth were soon to be recovered in Antarctica. An ad hoc committee was convened on 11 No- vember 1977 in Washington, D.C. The meeting included representatives of NSF (Mort Turner), the fi eld party (Wil- liam Cassidy), the Smithsonian Institution (SI, Brian Ma- son of the Natural History Museum and Ursula Marvin of the Astrophysical Observatory), National Aeronautics and Space Administration (NASA, Don Bogard of John- son Space Center and Bevan French of NASA headquar- ters), and the scientifi c community (including Jim Papike) (Antarctic Meteorite Working Group, 1978). This meeting produced ?a plan for the collection, processing, and dis- tribution of the U.S. portion of the Antarctic meteorites collected during 1977-78? (Antarctic Meteorite Working Group, 1978:13). However, much of the groundwork for this system of interagency cooperation (which ultimately was formalized as the three-agency agreement between NASA, NSF, and SI) and distribution of samples was laid before the meeting. Brian Mason (Smithsonian Institution, personal communication, 2004) recounted a conversation with Mort Turner where the opinion expressed was that meteorites collected by U.S. fi eld expeditions should prop- erly become U.S. government property. It was agreed that NASA would provide short-term curation modeled on, but less rigorous than, standards for lunar rock curation, while the Smithsonian would assume responsibility for classifi cation and long-term curation and storage. The collection effort evolved into what is now known as the Antarctic Search for Meteorites (ANSMET). Thirty full seasons have now been completed with the recovery of more than 16,000 meteorites? more than were col- lected over the entire Earth in the previous 500 years. The fi eld party grew from three members initially, with six to eight members during much of its history, and peaked at 12 members split between two fi eld parties, with one supported by NASA with the specifi c objective of increas- ing the collection of martian meteorites. The NSF Divi- sion of Polar Programs, with decades of experience in exploring the harsh Antarctic environment, provides sup- port for the ANSMET. Currently, the ANSMET program is run by Ralph Harvey, an associate professor at Case Western Reserve University. Each year, teams of four to eight scientists work together collecting meteorites in re- mote fi eld locations for about six weeks during the aus- tral summer (November? January). Their primary goal is to recover a complete and uncontaminated sampling of meteorites. Systematic searches are conducted as a series of 30-m-wide parallel transects by snowmobile on areas of snow-free blue ice. If the concentration is high, transects by snowmobile are replaced by searching on foot, ensur- ing the recovery of meteorites as small as 1 cm in diameter. Many stranding surfaces are large enough to require sev- eral seasons in the same area. It is interesting to note that as the program evolved, the number of meteorites recovered changed dramatically. Starting with 11 meteorites in 1976, ANSMET averaged H11011200 meteorites per year from 1976 to 1984, before ramping up to an average of nearly 600 meteorites from 1985 to 2001. This average is remarkable given the cancel- lation of the 1989 fi eld season due to logistical problems and the intentional exploration of areas with greater and lesser numbers of meteorites to average out the curatorial workload from year to year. During 2002? 2006, an aver- age of more than 900 meteorites was recovered each year, including two seasons of 1200H11001 meteorites. The astounding success of the Antarctic meteorite pro- grams of the United States and Japan have spurred a number of other efforts, including those from Europe (EUROMET) (Folco et al., 2002) and China (Lin et al., 2002). Indeed, a few privately funded expeditions have actually recovered meteorites in Antarctica. These events caused the Antarctic Treaty Organization to encourage member countries to take measures to protect this valuable scientifi c resource. The U.S. government, through the NSF, responded by implementing a federal regulation (45 CFR 674; National Science Foun- dation, 2003) that codifi ed, for the fi rst time, collection and curatorial standards used by the U.S. Antarctic Meteorite Program. It is important to note that other national govern- ments and government consortia (e.g., EUROMET) adhere to similar standards, although each has standards adapted to their unique situation. SMITHSONIAN?S ROLE IN THE U.S. ANTARCTIC METEORITE PROGRAM While the Smithsonian?s role has primarily been in classifi cation and curation, it has been greatly strengthened by the participation of several SI staff in the fi eld efforts over the years (Figure 2). Ursula Marvin of the Smithso- nian Astrophysical Observatory, who played a pivotal role in both the initial formation and long-term manage- ment of the program over the next three decades, was the fi rst Smithsonian participant in 1978? 1979 and returned in 1981? 1982, joined by Bob Fudali of the Division of 31_McCoy_pg387-394_Poles.indd 38931_McCoy_pg387-394_Poles.indd 389 11/17/08 9:56:14 AM11/17/08 9:56:14 AM 390 SMITHSONIAN AT THE POLES / MCCOY, WELZENBACH, AND CORRIGAN Meteorites, a long-time associate of Cassidy. Subsequently, Fudali (1983? 1984, 1987? 1988), meteorite collection managers Twyla Thomas (1985? 1986) and Linda Welzen- bach (2002? 2003, 2006? 2007), and postdoctoral fellows Sara Russell (1996? 1997) and Cari Corrigan (2004? 2005) served on the ANSMET fi eld parties. While the collection effort was shared by many, the classifi cation of Antarctic meteorites has been largely the responsibility of two individuals, Brian Mason and Tim McCoy. Mason volunteered his services during the forma- tive stages of the program, and it would be hard to have found a more perfect individual to undertake the challenge of classifying thousands of individual meteorites. During his tenure at the American Museum of Natural History in New York and subsequently at the Smithsonian Institu- tion, Mason had examined virtually every type of mete- orite known, pioneered the use of mineralogical data in the classifi cation of meteorites in his seminal 1963 paper (Mason, 1963), and, when faced with an unusual Antarc- tic meteorite, could quickly recall other similar meteorites he had examined during his long career. During his long tenure with the ANSMET program, Mason would go on to classify more than 10,000 individual meteorites, includ- ing a considerable number of Japanese meteorites during a visit to the National Institute of Polar Research in 1982. McCoy was hired in large part because of his experience and interest in classifi cation of and research on Antarctic meteorites and has overseen classifi cation, with the help of Welzenbach and a cadre of students and postdoctoral fellows, of more than 5,000 meteorites. After macroscopic descriptions are completed at NASA?s Johnson Space Center, a small chip is sent to the Smithsonian for classifi cation. In the earliest days of the program, a thin section was prepared for every meteorite, and mineral compositions were measured using the elec- tron microprobe. As the numbers of meteorites ramped up between 1984 and 1988, it became clear that this la- borious, time-consuming technique was producing an unacceptably large backlog of meteorites awaiting clas- sifi cation. Mason saw a need for a quicker technique to separate and classify the myriad of equilibrated ordinary chondrites. In 1987, he returned to a technique he had successfully applied in the 1950s and early 1960s? oil im- mersion. The rapid determination of the composition of a few olivine grains from each meteorite then became and remains the method by which 80%? 90% of all U.S. Ant- arctic meteorites are classifi ed. Unequilibrated ordinary, carbonaceous, and enstatite chondrites and achondrites are sent for thin section preparation, along with some meteorites that cannot be confi dently classifi ed due to brecciation, shock, or severe weathering. The Smithsonian?s Antarctic thin-section li- brary now contains over 5,000 thin sections, and H11011200 new sections are prepared each year. Mineral composi- tions (olivine and orthopyroxene for most chondrites; olivine, pyroxene, and plagioclase for achondrites) are determined using the JEOL JXA-8900R electron micro- probe. The Smithsonian prepares brief descriptions, tables of data, and digital petrographic images that are published in the Antarctic Meteorite Newsletter (Satter- white and Righter, 2006), which is also posted on the Web. Antarctic iron meteorites, which are found at very modest rates, are permanently transferred. The Smithson- ian has unique capabilities for processing iron meteorites and handles all processing, curatorial, and classifi cation of irons. For more than 30 years, the responsibility for FIGURE 2. Linda Welzenbach collects an achondrite meteorite at Larkman Nunatak during the 2006? 2007 ANSMET fi eld season. 31_McCoy_pg387-394_Poles.indd 39031_McCoy_pg387-394_Poles.indd 390 11/17/08 9:56:15 AM11/17/08 9:56:15 AM U.S. ANTARCTIC METEORITE PROGRAM 391 the description and curation of Antarctic meteorites fell to Roy S. Clarke Jr., whose specialty in the metallogra- phy of iron meteorites and oversight of the non-Antarctic collection made him an ideal choice. While all meteorites are classifi ed, the Smithsonian?s major task is identifying those specimens that are of particular interest to scien- tists and that would be worthy of further study. Figure 3 illustrates the results of these efforts, indicating the num- ber of samples recovered, the total number of meteorites, and the subset of meteorites that are not equilibrated or- dinary chondrites. The number of meteorites recovered increased steadily from an average of H11011300 (1977? 1984) to greater than 1,000 per year (2002? 2006) as collecting techniques improved and fi eld parties grew in size and number. The number of samples was sometimes greater than the number of meteorites due to the collection of a small number of terrestrial rocks mistaken for mete- orites. The number of the most scientifi cally interesting specimens, those other than equilibrated ordinary chon- drites (e.g., unequilibrated ordinary chondrites, carbona- ceous and enstatite chondrites, achondrites, and irons), remained constant at H1101150/year. (The sharp dip in 1989 was due to a cancelled fi eld season.) The apparent discon- nect between the number collected and those of greatest scientifi c interest is due to the occurrence of meteorites that break up in the atmosphere and possibly shower lo- cal areas with thousands of individual fragments. While most scientifi c studies focus on the small subset of the most interesting specimens, the collection as a whole still offers clues to ice movements related to concentration mechanism and the infl ux of meteoritic material to Earth over time (Harvey, 2003). Only through systematic col- lection and classifi cation of all the meteorites can these latter studies be undertaken. The other major obligation of the SI in the U.S. Ant- arctic Meteorite Program was serving as the long-term cu- ratorial facility for specimens (Figure 4). In 1983, the SI opened its Museum Support Center in Suitland, Maryland. This state-of-the-art collections facility is centered on four pods (football-fi eld-sized buildings H1101150 feet (H1101115 m) high) FIGURE 3. The number of samples and recovered meteorites and those meteorites that are not equilibrated ordinary chondrites (EOCs) from 1977 to 2007. 31_McCoy_pg387-394_Poles.indd 39131_McCoy_pg387-394_Poles.indd 391 11/17/08 9:56:16 AM11/17/08 9:56:16 AM 392 SMITHSONIAN AT THE POLES / MCCOY, WELZENBACH, AND CORRIGAN connected by a corridor of offi ces and laboratories. Shortly after this facility opened, planning began for building what became essentially a duplicate of the dry nitrogen storage facility for Antarctic meteorites at Johnson Space Center in Houston, and the new museum storage facility opened in the fall of 1986. The fi rst signifi cant transfer (126 speci- mens) of Antarctic meteorites to the Smithsonian occurred in 1987. Regular annual transfers from Johnson Space Cen- ter to the museum began in 1992, and the fl ow of meteor- ites increased tremendously in 1998. At that point, the Me- teorite Processing Laboratory at Johnson Space Center was essentially full, and the subsequent infl ux of newly recov- ered meteorites necessitated the transfer of large numbers of specimens to the SI. By the end of 2004, more than 11,300 individual specimens had been transferred to the museum. When coupled with the chips and thin sections used for the initial classifi cation, Antarctic meteorites now represent more than 80% of named meteorites in the Smithsonian collection and more than 70% of all specimens. These per- centages alone demonstrate the spectacular impact of the Antarctic Meteorite Program on the Smithsonian?s meteor- ite collection. During the 30 years of the U.S. Antarctic Meteorite Program, Smithsonian personnel have fulfi lled a num- ber of other roles. The program is managed by a three- member Meteorite Steering Group with representatives from NASA, NSF, and the Smithsonian. Recommendations on sample allocations are made by the Meteorite Working Group, a 10-member panel that also includes members of the academic and research communities. Smithsonian per- sonnel from both the Natural History Museum and the Astrophysical Observatory have actively participated or FIGURE 4. Meteorite storage laboratories at the Smithsonian Museum Support Center in Suitland, Maryland, modeled on the facility used for lunar rocks at NASA?s Johnson Space Center. The water- and oxygen-free nitrogen gas in the cabinets keeps meteorites from oxidizing and free from contamination by environmental pollutants such as organic compounds, heavy metals, and salts, which could reduce the scientifi c value of the specimens. Photo by Chip Clarke, SI. 31_McCoy_pg387-394_Poles.indd 39231_McCoy_pg387-394_Poles.indd 392 11/17/08 9:56:23 AM11/17/08 9:56:23 AM U.S. ANTARCTIC METEORITE PROGRAM 393 led these committees throughout the history of the pro- gram. Additionally, the Smithsonian provides selected samples of Antarctic meteorites for exhibits throughout the world, including meteorites on display at the Crary Science and Engineering Center in McMurdo Station, per- haps the southernmost display of Smithsonian objects. SCIENTIFIC VALUE OF ANTARCTIC METEORITES While the U.S. Antarctic Meteorite Program has had a dramatic infl uence on the size of the Smithsonian mete- orite collection, it is the information that these priceless samples hold that is of greatest benefi t. While listing the full range of scientifi c discoveries is beyond the scope of this paper, we list a few examples. Brian Mason and Smithsonian volcanologist Bill Melson published the fi rst book-length treatise on Apollo 11 samples in 1970 (Mason and Melson, 1970), and Mason remained involved in the study of lunar samples through the end of the Apollo Program. In 1982, Mason described the Antarctic meteorite ALH A81005 as con- taining clasts that ?resemble the anorthositic clasts de- scribed from lunar rocks? (Mason, 1983). From his earlier work, Mason knew this was the fi rst lunar meteorite but presented his fi ndings in a typically understated manner so as not to undercut the considerable research that would be forthcoming. Today, we recognize more than three dozen distinct lunar meteorites. A remarkable feature of these meteorites is that they commonly exhibit very low abun- dances of the radioactive element thorium. In contrast, the area sampled by the Apollo missions on the equato- rial near side of the Moon typically has elevated thorium concentrations indicative of the thin crust and extensive volcanism that occurred in that region. For this reason, lunar meteorites are thought to represent a much broader representative sampling of the lunar surface and are the subject of intense scrutiny as the U.S. plans for a return to the Moon (Korotev et al., 2003). The realization that lunar meteorites had been launched by impacts from the surface of the Moon and escaped the heating that many predicted would melt them completely reinvigorated debate about whether certain meteorites ac- tually originated on Mars. This debate was largely settled in 1981, when Don Bogard and colleagues at Johnson Space Center showed that gases trapped inside impact melt pockets in the Antarctic meteorite EET A79001 matched those measured in the martian atmosphere by the Viking lander. These samples, now numbering several dozen, pro- vide the only materials from Mars that we have in our lab- oratories. Although they lack geologic context, study of these rocks has posed many of the questions driving Mars exploration. This was never more true than when McKay et al. (1996) argued that ALH 84001 contained evidence of past microbial life in the form of distinctive chemical, mineralogical, and morphological features. Although the result has been vigorously debated for over a decade, it is clear that this single paper reinvigorated NASA?s Mars Exploration Program. The founding of the NASA Astro- biology Institute and the launch of the Mars Exploration Rovers Spirit and Opportunity, which continue to oper- ate after three years on the surface of Mars and on which the senior author is a team member, were spurred in part by the idea that ancient life may have existed on Mars. It is truly remarkable that a modest program of collecting meteorites? the poor man?s space probe? prompted the initiation of major research and spacecraft efforts! Among the signifi cant advances in meteoritics within the last decade, one of the most noteworthy is the rec- ognition of meteorites that are intermediate between the primitive chondrites formed as sediments from the solar nebula and achondrites that sample differentiated bodies with cores, mantles, and crusts, like Earth. These meteor- ites, termed primitive achondrites, experienced only par- tial melting and differentiation, after which the process was halted. These meteorites may offer our best clues to how our own planet differentiated. While such meteorites have been known for more than a century, they were few in number and largely viewed as curiosities. The vast num- bers of meteorites recovered from Antarctica have pushed these meteorites into prominence, as major groupings have emerged. Among these meteorites, one is truly remark- able. Graves Nunatak (GRA) 95209 contains metal veins that sample the earliest melting of an asteroid as it began to heat up more than 4.5 billion years ago ( McCoy et al., 2006). These veins record a complex history of melting, melt migration, oxidation-reduction reactions, intrusion into cooler regions, cooling, and crystallization. The single most remarkable feature of this meteorite is the presence of millimeter-size metal grains that contain up to a dozen graphite rosettes tens of micrometers in diameter. Within a single metal vein, carbon isotopic compositions (H9254 13 C) can range from H1100250 to H1100180?. These graphite grains formed not in the parent asteroid during melting but during nebu- lar reactions. This isotopic heterogeneity is even more re- markable when we consider that a single millimeter-sized metal grain within this meteorite has a greater carbon iso- topic heterogeneity than all the natural materials on Earth. Despite extensive heating, this asteroid did not achieve a 31_McCoy_pg387-394_Poles.indd 39331_McCoy_pg387-394_Poles.indd 393 11/17/08 9:56:27 AM11/17/08 9:56:27 AM 394 SMITHSONIAN AT THE POLES / MCCOY, WELZENBACH, AND CORRIGAN homogeneous carbon isotopic composition. In a very real way, this sample gives us a glimpse into the processes that occurred in the solar nebula during the birth of our solar system as well during the heating, melting, and differentia- tion of our Earth. CONCLUSIONS The meteorite collection and the insights provided by the infl ux of a tremendous number of Antarctic meteorites continue to enrich the Smithsonian collections and offer opportunities for research among its staff. Much of the current emphasis is in the burgeoning fi eld of astrobiology. Understanding the fate of carbon during the differentiation of planets forms another link in understanding this funda- mental element from its birth in other stars to the role it plays today in biologic evolution. Scientists at the Smith- sonian use these meteorites to ask fundamental questions that they then set about answering through participation in spacecraft missions to the Moon, Mars, asteroids, and comets. Rather than supplanting meteorites as a major source of information, samples returned from these bodies will make Antarctic meteorites more scientifi cally valuable as they continue to provide the framework as we continue to ask and answer the questions of our solar system?s birth, evolution, and destiny. ACKNOWLEDGMENTS This paper reports the results of an effort shared by hundreds of individuals over three decades. These include the members of the meteorite search teams, the curatorial teams at Johnson Space Center and the Smithsonian Insti- tution, the administrative support at NASA, the National Science Foundation, and the Smithsonian, and the efforts of thousands of scientists who have studied Antarctic me- teorites. Among these, Don Bogard (NASA Johnson Space Center), Ursula Marvin (Smithsonian Astrophysical Ob- servatory), and Bill Cassidy (University of Pittsburgh) have given us insights over the years into the formative stages of the U.S. Antarctic Meteorite Program. Our current partners in the curatorial and collection efforts Ralph Harvey (Case Western Reserve University), John Schutt (ANSMET), Kevin Righter (NASA Johnson Space Center), and Cecilia Satterwhite (Jacobs) have been particularly helpful, as have our colleagues at the Smithsonian, Brian Mason, Roy S. Clarke Jr., and the late Gene Jarosewich, who preceded us in these efforts and have been unfailingly supportive. We dedicate this paper to our colleague Gene Jarosewich, who never wavered in his enthusiasm for meteorites. LITERATURE CITED Antarctic Meteorite Working Group. 1978. Antarctic Meteorite News- letter, No. 1(1). http:// www-curator .jsc .nasa .gov/ antmet/ amn/ previous_ newsletters/ ANTARTIC_ METERORITE_NEWSLETTER_ VOL_ 1_ NUMBER_ 1 .pdf (accessed 15 August 2008). Cassidy, W. A. 2003. Meteorites, Ice, and Antarctica: A Personal Ac- count. Cambridge: Cambridge University Press. Folco, L., A. Capra, M. Chiappini, M. Frezzotti, M. Mellini, and I. E. Tabacco. 2002. The Frontier Mountain Meteorite Trap. Meteoritics and Planetary Science, 37: 209? 228. Harvey, R. 2003. The Origin and Signifi cance of Antarctic Meteorites. Chemie der Erde, 63: 93? 147. Korotev, R. L., B. L. Jolliff, R. A. Zeigler, J. J. Gillis, and L. A. Haskin. 2003. Feldspathic Lunar Meteorites and Their Implications for Compositional Remote Sensing of the Lunar Surface and the Com- position of the Lunar Crust. Geochimica et Cosmochimica Acta, 67: 4895? 4923. Lin, C.-Y., F. S. Zhang, H.-N. Wang, R.-C. Wang, and W.-L. Zhang. 2002. ?Antarctic GRV9927: A New Member of SNC Meteorites.? Thirty-third Annual Lunar and Planetary Science Conference, March 11? 15, Lunar and Planetary Institute, Houston, Tex. Mason, B. H. 1963. Olivine Composition in Chondrites. Geochimica et Cosmochimica Acta, 27: 1011? 1023 ??? . 1983. Antarctic Meteorite Newsletter, No. 6 (1). http:// curator .jsc .nasa .gov/ antmet/ amn/ previous_ newsletters/ ANTARTIC_ METERORITE_ NEWSLETTER_ VOL_ 6_ NUMBER_ 1 .pdf (accessed 15 August 2008). Mason, B. H., and W. G. Melson. 1970. The Lunar Rocks. New York: Wiley-Interscience. McCoy, T. J., W. D. Carlson, L. R. Nittler, R. M. Stroud, D. D. Bogard, and D. H. Garrison. 2006. Graves Nunataks 95209: A Snapshot of Metal Segregation and Core Formation. Geochimica et Cosmochi- mica Acta, 70: 516? 531. McKay, D. S., E. K. Gibson, K. L. Thomas-Keprta, H. Vali, C. S. Romanek, S. J. Clemett, X. D. F. Chiller, C. R. Maechling, and R. N. Zare. 1996. Search for Past Life on Mars: Possible Relic Biogenic Activity in Martian Meteorite ALH84001. Science, 273: 924? 930. National Science Foundation. 2003. U.S. Regulations Governing Ant- arctic Meteorites. 45 CFR Part 674. http:// www .nsf .gov/ od/ opp/ antarct/ meteorite_ regs .jsp (accessed 15 August 2008). Satterwhite, C., and K. Righter, eds. 2006. Antarctic Meteorite News- letter. http:// curator .jsc .nasa .gov/ antmet/ amn/ amn .cfm (accessed 15 August 2008). Shima, M., and M. Shima. 1973. Mineralogical and Chemical Composi- tion of New Antarctica Meteorites. Meteoritics, 8: 439? 440. Welten, K. C., C. Alderliesten, K. Van der Borg, L. Lindner, T. Loeken, and L. Schultz. 1997. Lewis Cliff 86360: An Antarctic L-chondrite with a terrestrial age of 2.35 million years. Meteoritics and Plan- etary Science, 32: 775? 780. 31_McCoy_pg387-394_Poles.indd 39431_McCoy_pg387-394_Poles.indd 394 11/17/08 9:56:27 AM11/17/08 9:56:27 AM Index AAUS. See American Academy of Underwater Sciences Adams, David Hempelman, 53?54 Advanced Microwave Scanning Radiometer-EOS, 311 Advanced very high resolution radiometer (AVHRR), 310 Ahlmann, Hans W., 26 Ahmaogak, George, Sr., 104 Ahwinona, Jacob, 100, 105 Aiken, Martha, 100 Airy, George B., 17 Alaska. See also Barrow, Alaska; Biological invasion; Exhibits; Native peoples; Whaling art, public discovery, 67?70 ethnological collecting, 62?65, 89?96, 99?100, 118?121, 131?132 ethnological exhibits, 65?66 Gambell, and SIKU, 134?138 Smithsonian connection, revitalizing, 70?72 southwest (Yup?ik homeland), 80, 81 St. Lawrence Island, and SIKU, 134?138 Aleut people. See Native peoples Alutiiq people. See Native peoples American Academy of Underwater Sciences (AAUS), 242 Amundsen-Scott South Pole Station, 382 Anchorage Museum of History and Art, xiv, 71, 80, 85 Andersson, Berit, 51 Andr?e, Solomon August, 50?51, 53 ballooning, 50?51 cause of death, 52, 53 expedition camp, 51?52 archaeologic study, 50?54, 59 soil studies, 52?54 Andrew, Frank, 82?84, 85, 86 Andrew, Nick, 87 Annals of the IGY (1959), 31 Annual Reports of the Bureau of Ethnography, xiii 32_Index_pg395-405_Poles.indd 39532_Index_pg395-405_Poles.indd 395 11/17/08 10:32:58 AM11/17/08 10:32:58 AM 396 INDEX Antarctic Circumpolar Current copepod distribution, 153 nonpelagic development, 185?187, 192 productivity, 312 Antarctic Continental Shelf, isolation and speciation, 185?187 Antarctic Convergence, 145, 147 Antarctic deep-sea benthic biodiversity (ANDEEP), 191 Antarctic Marine Ecosystem Ice Edge Zone (AMERIEZ), 290 Antarctic Marine Living Resources (AMLR), 289 Antarctic Meteorite Working Group, 389, 392?393 ?Antarctic Research?A European Network for Astronomy? (ARENA), 385 Antarctic Search for Meteorites (ANSMET), 389, 390 Antarctic Submillimeter Telescope and Remote Observatory (AST/RO), x, xiv, 361, 362, 363, 369?372, 381?384 Antarctic Treaty, 23 Antarctic Treaty Organization, 389 Antarctica. See also Biological invasion; Cosmology, Antarctica East Base (Stonington Island), 50, 54?55, 56, 59 Marble Point, 50, 56?59 West Base, 54 Anthropology, 115?126 circumpolar, 70 cultural, 64 foundations, 116?121 IPY goal, 121 museum, 62 physical, 66?67 salvage, 118 theory of, 65?66 Apangalook, Leonard, Sr., 134?138 Aporta, Claudio, 134 Applegate, S., 65 Archaeology. 115?126. See also Andr?e, Solomon August; Antarctica circumpolar, 70 historic, 49?60 impact on Smithsonian, 66?67 tourism, 60 Arcminute Cosmology Bolometer Array Receiver (ACBAR), 363 Arctic as cultural construct, 115 differing meanings, 115?117 as homeland, 116 as ?last imaginary place,? 115 Arctic studies, 61?74, 131?132. See also International Polar Year climate change, 65, 71, 72?74 hardships, 18?19 public interest, 49 Arctic Studies Center (ASC), x, xiv, 70, 133?134 Army Signal Corps, xiii, 5, 16, 19, 63 Arutiunov, Sergei, 70 Assault on the Unknown (Sullivan), 31 AST/RO. See Antarctic Submillimeter Telescope and Remote Observatory Aurora(s), 3, 7 Background Imaging of Cosmic Extragalactic Polarization (BICEP), 364 Bacterioplankton, productivity, 299?306 Baffi n Island, Boas visit, 117?118, 131 Baird, Spencer F. anthropology, 118, 119 arctic studies, 62?64, 65, 131 international cooperation, 17 IPY-1, xiii, 94 Polaris Expedition (1871), 18?19 Ballooning expeditions. See Adams, David Hempelman; Andr?e, Solomon August Balloon Observations of Millimetric Extragalactic Radiation and Geophysics (BOOMERANG), 361 Barrow, Alaska, 89?96 as archaeological site, 60 early expeditions, 90?92 ethnological collecting, 64?65, 89?96, 131 geography and climate, 89 as IPY-1 station, xiii, 5, 6, 19, 64?65, 89, 92?95, 119 map, 91 post-IPY research, 95?96 whaling, 99?111 Bartlett Inlet, 267 Bearded seals. See Seals Beckman, Arnold, 28 Beechey, Frederick W., 90 Beluga whales. See Whales Bennett, Jim, 36 Benthic communities, link with pelagic, 218?219 Berkner, Lloyd, 9, 24?28, 29, 31, 35 Bertrand, Kenneth, 18 Bessels, Emil, 19 Biological invasion, 347?355 climate change, 351?354, 355 commercial shipping, 349?350, 352, 353?354 crabs, 348?349, 350?351 currents, 351, 352?353 detection and measurement, 355 fi sheries, 354 human-mediated, 347?348, 349?350 live trade, 349 oil and gas industry, 349?350, 353?354 prevention, 355 propagule supply, 349?350 resistance, 350?351 shoreline development, 353?354 species, Alaska, 348 species, Antarctic, 348?349 temperate-polar pattern, 348?349 32_Index_pg395-405_Poles.indd 39632_Index_pg395-405_Poles.indd 396 11/17/08 10:32:59 AM11/17/08 10:32:59 AM INDEX 397 temperature, 350?355 tourism, 354 Black holes, observations. See Antarctic Submillimeter Telescope and Remote Observatory Black, Richard, 54 Blee, Catherine, 55 Boas, Franz, 64?65, 70, 117?118, 131 Bockstoce, John, 91?92 Bodenhorn, Barbara, 96 Bogard, Don, 389, 393 Bowhead whales. See Whales Bowman, Isaiah, 6, 8 Brine channels, 244 Britannic Challenger (balloon), 53?54 British Association for the Advancement of Science, 14?15, 16 Brooding species. See Bryozoan, Antarctica; Invertebrates, marine Brouger-Lambert Law, 273 Brower, Jane, 100, 110 Brower, Ronald, Sr., 100, 102, 104?108, 110 Bruce, Miner, 100 Bryozoans, Antarctic, 205?219 Budd, William, 19 Bulletin of International Simultaneous Observations, 4?5, 16 Buoyancy compensator (BC), 246?247 Bureau of American Ethnology, 65, 94, 118 Bush, Vannevar, 25?26 Byrd, Richard E., 54 Calista Elders Council (CEC), 80, 82, 85 Canineq collection, 82 Carbon monoxide, as interstellar tracer, 369?372, 373, 381?384 Caribou hunting, 95, 101, 121 Cassidy, William, 388?390 Cawood, John, 15, 19?20 CDOM. See Chromophoric dissolved organic matter The Central Eskimo (Boas), 64?65 Chapman, Sydney, 1, 9, 24?28, 31, 35 Chaussonnet, Val?rie, 70 Chlorophyll CDOM cycling, 321?322, 326?329 kelp productivity, 272, 273, 276, 277 phytoplankton and bacteria, 302 Southern Ocean, productivity, 310?311 Christie, Samuel Hunt, 15 Chromophoric dissolved organic matter (CDOM), 319?331. See also Chlorophyll; Pack ice absorption spectra, 322?331 ecosystem effects, 320 photobleaching, 320 sources, 320, 330 storage test, 323?324 study site, 321?322 transect studies, 324 Clarke, Roy S., Jr., 390?391 Climate change. See also Biological invasion; Climate studies arctic sites, loss of, 59 arctic storms, 136?137 arctic studies, 26, 72?73 birds, 138 bryozoans, 218 indigenous observations, 129?139 krill, 286, 287, 294?295 marine mammals, habitat and behavior, 137?138 sea ice, 134?138 Stonington Island, 56, 59 Climate studies, 1?11. See also Arctic studies; Cooperation, scientifi c Clouds, interstellar AST/RO observations, 363, 369?372, 381?384 HEAT observations, 373?379 life cycle, 373?374 Coastal Zone Color Scanner (CZCS), 310 Cold adaptation, 253?262 Collaboration. See Cooperation, scientifi c Collins, Henry B., 67, 69, 132 Comit? Sp?cial de l?Ann?e G?ophysique Internationale (CSAGI), 9, 27, 32?33 Cooperation, scientifi c, 13?20. See also International Geophysical Year astronomy, 16?17 Brussels meeting (1853), 19 Global Atmospheric Research Program (GARP), 9?10 Global Weather Experiment (GWE), 9?10 IPY-1, 13?20, 118 Magnetic Crusade, 14?15, 19 climate studies, 15?16 Venus, transits of, 14, 17?18, 19 Copepods, pelagic calanoid. See Pelagic calanoid copepods Corrigan, Cari, 390 Cosmic Background Explorer (COBE), 360 Cosmic microwave background (CMB) radiation, 360?364 Cosmology, Antarctica, 359?364 atmospheric conditions, 360?362 development, 360?361 as quantitative science, 359?360 site testing, 361?362 telescopes and instruments, 363?364 Crustaceans. See Southern Ocean; Biological invasion Crary Science and Engineering Center, 393 Crowell, Aron, 70, 71, 72?74, 82 CSAGI. See Comit? Sp?cial de l?Ann?e G?ophysique Internationale Currents. See Antarctic Circumpolar Current; Biological invasion 32_Index_pg395-405_Poles.indd 39732_Index_pg395-405_Poles.indd 397 11/17/08 10:33:00 AM11/17/08 10:33:00 AM 398 INDEX Dall, William Healy, 18, 63, 64 ?Dark Sector,? 361. See also Cosmology, Antarctica Darlington, Jennie, 54 Davis, J. E., 17 Davis Strait, 6 ?Deal 1? payload, 41?42 Defense Meteorological Satellite Program, 311 Dirksen, Everett, 28?29 Dissolved organic matter (DOM), 319, 330. See also Chromophoric dissolved organic matter Diving, x, xiv, 241?251 access through sea ice, 244?245, 267 animal and bird physiology, 265?270 animals as hazard, 248?249 decompression, 251 ?diver down? fl ag, 248 emergencies, 249?250 environment, 243?244 environmental protection, 250 equipment, 245?247 fast ice, 243, 244, 265?266 ice-edge, 248 ice formation, 243?244 krill studies, 290?295 McMurdo Sound, 265?267, 270 minimum qualifi cation criteria, 242 pack ice, 244, 248, 290?292 physiological considerations, 250?251 polar operations, 244?247 science, necessity, 253?254, 261?262 ?suit squeeze,? 247 underwater visibility, 244, 248 weight and trim systems, 246?247 Dome A, astronomical observations from, 377?378, 384?385. See also High Elevation Antarctic Terahertz Telescope Dome C, astronomical observations from, 385, 386 Doubly labeled water (DLW) technique, 339 Dowdeswell, Julian, 18 Dragovan, Mark, 361 Drake Passage, 186?187, 192 Dufek, George J., 56?57 Eagle (balloon), 50?51. See also Andr?e, Solomon August Eicken, Hajo, 134 Ekholm, Nils, 51 Elephant Island. See Octopods, abundance Ellesmere Island, astronomical observations from, 385 Elliott, Henry W., 63 Elson, Thomas, 90 Emission Millimetrique (EMILIE), 360?361 Emmons, George T., 100, 106 Emperor penguins, 265?270 Environmental change. See Climate change Environmental cleanup, at Antarctic bases, 50, 54?55 Eskimos. See Native peoples; Yupik Eskimos; Yup?ik Eskimos The Eskimos About Bering Strait (Nelson), 65 Espy, James P., 3 Ethnological Results of the Point Barrow Expedition (Murdoch), 64?65, 110?111 Ethnology. See also Alaska; International Polar Year; Murdoch, John; Native peoples; Smithsonian Institution; Turner, Lucien early Smithsonian efforts, 62?65, 89?96, 99?100, 116? 121, 131?132 early Smithsonian exhibits, 65?66 public discovery of Eskimo art, 67?70 sale or barter of items, 94, 101 Smithsonian?Alaska connection, revitalizing, 70?72 Euphausia superba. See Krill, Antarctic Evolutionary temperature adaptation, 183 Ewing, Heather, 62 Exhibits Agayuliyararput/Our Way of Making Prayer, 79?80, 81?82 Arctic: A Friend Acting Strangely, x, xii, 73, 130 Crossroads Alaska, 70?71 Crossroads of Continents: Cultures of Siberia and Alaska, 69?70, 122 Crossroads Siberia, 70 The Far North: 2000 Years of American Indian and Eskimo Art, 67?68, 122 Inua: Spirit World of the Bering Sea Eskimo, 68?69, 70, 79?80, 122 Looking Both Ways: Heritage and Identity of the Alutiiq People, 71 Our Universes, 82 The People of Whaling, 100 Sharing Knowledge: The Smithsonian Alaska Collections, 72, 82, 100 Vikings: The North Atlantic Saga, 70 Expeditions. See also specifi c vessels; Barrow, Alaska; Henry, Joseph Austro-Hungarian North Pole Expedition (1872), 5 Belgica expedition, collections, 144, 206 Challenger expedition (1873?1876), 144, 181, 187, 189 Eclipse Expedition, 119 German Antarctic Expedition, 54 Norwegian-British-Swedish Antarctic Expedition (1949?1952), 26 Polaris Expedition (1871), 18?19 Ronne Antarctic Research Expedition (RARE), 55 Second Byrd Antarctic Expedition (1933?1935), 8 Field, Cyrus, 17 Fisher, William J., 65 32_Index_pg395-405_Poles.indd 39832_Index_pg395-405_Poles.indd 398 11/17/08 10:33:02 AM11/17/08 10:33:02 AM INDEX 399 Fishing. See also Biological invasion bryozoan abundance, 218 octopod abundance, 197?202 Fitzhugh, William, x, xi, 68?70, 73?74, 79 Fort Conger, Lady Franklin Bay station, xiii, 5, 60, 65, 131 Fraenkel, Knut, 51, 53 Franklin, Sir John, 91 search expeditions, 91?92, 116 Fraser, Ronald, 28 French, Bevan, 389 Friedman, Herbert, 45?46 Fudali, Bob, 389?390 Gauss, Carl Friedrich, 14 Georgi, Johannes, 7 German Polar Commission, 117 Giant molecular clouds (GMCs), 374, 375 Golovnev, Andrei, 70 G?ttingen Magnetic Union/Verein, 2, 14?15 Gray whales. See Whales Greely, Adolphus W., 5?6, 19, 65, 132 Greenland, astronomical observations from, 385 Hagen, John P., 39 Hall, Charles Francis, 18?19, 116?117 Hamilton, Joan, 84, 87 Harvey, Ralph, 389 Hayes, I. I., 18 HEAT. See High Elevation Antarctic Terahertz Telescope Heinzel, Karl, 41 Henry, Alfred J., 6 Henry, Joseph arctic studies, 62?64, 74 international cooperation, 14, 15, 17 Meteorological Project, 3, 15?16 Polaris Expedition (1871), 18?19 Herendeen, Edward Perry, 92?93 Her Many Horses, Emil, 82 Herschel, John, 2, 15 High Elevation Antarctic Terahertz (HEAT) Telescope, 373? 379, 385. See also PreHEAT High-resolution spectroscopic imaging, 375 Hillary, Edmund, 59 H.M.S. Plover, 91?92 Holm, Bill, 70 Holmes, William Henry, 65 Hrdlic?ka, Ale?, 66?67, 69 Hudson?s Bay Company, 15, 62, 90, 119 Hudson?s Bay Territory, 62, 119 Humboldt, Alexander von, 5, 14 Hunting. See also Caribou hunting; Whaling climate change, 135?138 endangered perspective on past, 122?123 Ice. See also Diving; Pack ice; Sea ice camps, 291?293 congelation, 243 fall formation, new patterns, 135 indigenous observations, 133?138 krill habitat, 286?295 pancake, 243 platelet, 243?244, 249 stalactites, 244 winter weather changes, 135?137 IGY. See International Geophysical Year International Astronomy Union (IAU), 26, 29 International cooperation. See Cooperation, scientifi c International Geophysical Year (1957?1958), xiii?xix, 1?2, 8?10, 23?31, 35?36 accomplishments, 9, 23 agenda-setting for, 24, 25?28, 31, 35 Berlin Congress (1878), 24 bryozoan collecting, 206 glaciology, 23, 26, 31 logo, 9 NASM collection, 37?45 organizing committee, 9, 27, 32?33 policy process, 23?33 political context, 24?25, 35, 59 publications, 31 purpose, 9 satellites, 9, 23, 24, 36 Soviet participation, 29?31, 36 World Data Centers, 9, 23, 29?31 International Polar Year, fi rst (1882?1883), xiii, 1, 5?6, 11 accomplishments, 6 ethnological collecting, 64?65, 89?96, 99?100, 118?121, 131 Humboldtean science, 5 international cooperation, 13?20, 118 as landmark event, 13 limitations, 6 political context, 24 reanalysis and reevaluation, 6 scientifi c vs. natural history agenda, 118 Smithsonian participation, 61?62, 64?65, 89?96 social sciences, 131 stations, xiii, 5?6 U.S. participation, 19 Weyprecht?s principles, 5 International Polar Year, second (1932?1933), xiii?xiv, 1, 6?8 accomplishments, 8 political context, 24 radio technology, 7?8 social sciences, 131 32_Index_pg395-405_Poles.indd 39932_Index_pg395-405_Poles.indd 399 11/17/08 10:33:03 AM11/17/08 10:33:03 AM 400 INDEX International Polar Year, third. See International Geophysical Year International Polar Year, fourth (2007?2008), x, xi, xiv?xv, 10?11 Smithsonian participation, 72?74 technological support, 10?11 themes, 10 International Polar Year: Astronomy from the Poles (AstroPoles), 384 International Research and Exchanges Board (IREX), 70 International Sun-Earth Explorer (ISEE-3), 37 International Union of Pure and Applied Physics (IUPAP), 26 International Union of Radio Science (URSI), 26 International Whaling Commission (IWC), 102 Interstellar clouds. See Clouds, interstellar Interstellar medium (ISM), 373, 375 Inuit people. See Native peoples Inuit Sea Ice Use and Occupancy Project (ISIUOP), 134 I?upiat Eskimos. See Native peoples I?upiat Heritage Center, 100, 102?103 Invertebrates, marine. See also Cold adaptation; Nonpelagic development benthic, 181?192, 205?219 brooding species, 181?192 IPY. See International Polar Year Isolated hole protocol (IHP), 267 ?Jesup II,? 70 John, Agatha, 87 John, Mark, 85 John, Paul, 82, 84, 85 John, Peter, 87 Johnson Space Center, xiv, 390, 392, 393 Joinville Island. See Octopods, abundance Juniper-C missile, 43 Kamkoff, Willie, 82 Kane, Elisha Kent, 18, 62 Kaplan, Susan, 68?69, 79 Kashevarov, Aleksandr, 90?91 Kayaks, 83?84, 85, 122 Kelp, arctic (Laminaria solidungula), 271?283. See also Chlorophyll ammonium, 272, 273, 276, 277, 280 biomass, 272, 274?275 Brouger-Lambert Law, 273 elongation (growth), 274, 278, 281?282 light availability, 271?283 oil and gas industry, 349?350, 353?354 pH, 272, 274, 278, 281 storms (wind speed), 281?282 total suspended solids (TSS), 271?283 Kennicott, Robert, 18, 61, 62, 64, 74 Khlobystin, Leonid, 70 Kjellstr?m, Rolf, 51 Kramer, S. Paul, 28 Krill, Antarctic (Euphausia superba), 285?295. See also Climate change; Diving biomass, 286 distribution, 286 ecosystem role, 286?287 as foundation species, 285?287 growth rates, 292?295 as keystone species, 285 life history, 287?289 population dynamics, 293?295 process cruise studies, 291?292 sea ice dynamics, 285?295 Krupnik, Igor, x, xi, 70, 73?74, 94, 133?134 Lactation. See Seals; Whales Lady Franklin Bay station, xiii, 5, 60, 65, 131 Laidler, Gita, 134 Lambda-Cold Dark Matter, 360 Laminaria solidungula. See Kelp, arctic Lang, Michael A., x, xi, 242 Larsen A ice shelf, collapse, 185 Larsen B ice shelf, collapse, 11, 185 Lecithotrophy, 183, 184?185, 186, 188, 189, 190, 192 Lenz, Mary Jane, 81, 82 Lewis, C. D., 68 Light availability. See Photosynthetically active radiation Lipps, J. H., 55 Lloyd, Humphrey, 15 Loki-Dart, 38, 43?44, 45 Loomis, Elias, 3, 15 Loring, Stephen, 74 The Lost World of James Smithson (Ewing), 62 Ludwig, George H., 40?42 Lundstr?m, Sven, 51 MacKay, Charles, 65 Magnetic Crusade, 14, 19 Magnuson, Warren, 29 Maguire, Rochefort, 91?92 Marble Point, Antarctica. See Antarctica Mars Exploration Program, 393 Marsh, H. Richmond, 100, 108 Marvin, Ursula, 389?390 Mason, Brian, 389, 390, 393 Mason, Otis, 66 Mather, Elsie, 85, 86, 87 Maury, Matthew Fontaine, 16 McCoy, Timothy, ix, 390 McGhee, Robert, 115 McMurdo Sound, as diving study area, 265?270 32_Index_pg395-405_Poles.indd 40032_Index_pg395-405_Poles.indd 400 11/17/08 10:33:04 AM11/17/08 10:33:04 AM INDEX 401 McMurdo Station, xiv, 265?267, 393 runways, 59 Meade, Marie, 81, 83 Melson, Bill, 393 Merriam, C. Hart, 66 Meteorites, Antarctic, ix, xiv, 387?394 classifi cation, 390?391 concentration, 388 Graves Nunatak (GRA) 95209, 393?394 iron, 390?391 lunar, 393 Mars, 393 NASA program, ix, xiv, 389, 390, 393 NSF program, 388?389 terrestrial ages, 388 types, 387?388 Meteorological Project, 3, 15?16 Meteorology, 1?11 Bergen school, 7 disciplinary period, 6 historical precedents, 2?5 IGY and, 1?2 international cooperation, 15?16 IPY and, 1, 2, 5?8, 10?11 nuclear testing and, 9 Milky Way Antarctic observations of center, 363, 369?372 constructing template, 375 star formation in. See Star birth (formation) Moore, Riley, 132 Morgan, Lewis Henry, 62 Moses, Phillip, 84?85 Moubray Bay, Antarctica, 266?267 Minimal Orbital Unmanned Satellite, Earth (MOUSE), 38 Muir, John, 64 Murdoch, John acquisition of objects, 99?100 and Baird, 131 on contact with Eskimos, 92 ethnological studies, 64?65, 69, 70, 71, 93?96, 110 Museum preservation, 36?37. See also specifi c museums Museum Support Center (MSC), 82?83, 391?392 MV Polar Duke, 290 Myer, Albert J., 16 Narwhal, dentition, 223?240 NASA. See National Aeronautics and Space Administration NASA/NASM Transfer Agreement, 37 National Academy of Sciences, 23, 27 National Aeronautics and Space Act (1958), 37 National Aeronautics and Space Administration (NASA). See also Meteorites, Antarctic oceanographic studies, 311 relationship with NASM, 37 Smithsonian collaboration, xi National Air and Space Museum (NASM), xiii?xiv, 35?47 National Anthropological Archives, 132 National Gallery of Art, 67?68, 122 National Geographic Society, 55 National Museum of the American Indian (NMAI), 71 whaling collection, 100 Yup?ik visits, 79, 81 National Museum of the American Indian Act (1996), 121 National Museum of Natural History (NMNH), ix, x, xiv. See also Arctic Studies Center; Exhibits anthropology, 118 ethnological collection, 132 Repatriation Offi ce, 71 whaling collection, 100 Yup?ik visits, 79 National Science Foundation (NSF), ix?x, xi, xiv. See also Meteorites, Antarctic Center for Astrophysical Research, x, 361 U.S. Antarctic Diving Program, 241?251 Native American Graves Protection and Repatriation Act (1990), 121 Native peoples. See also Repatriation; Whaling; Yupik Eskimos; Yup?ik Eskimos Ainu people, 70 Aleut people, 64?65 Alutiiq people, 64?65, 71 art, public discovery, 67?70 colonial concepts, 116 community anthropology, 123?125 elders, museum collaboration, 80?84, 99?111 endangered perspective on past, 122?123 ethnological collections, 65?66, 89?96, 99?100 Innu people, 64?65, 119?121, 123?125, 131 Inuit people, 122?123, 64?65, 116?117, 122, 223?239, 123?125, 65, 119?121, 131 I?upiaq people, 99?111 I?upiat people, 67?70, 90?92, 65?66, 89?96, 95, 92?95, 92, 94, 99?100, 101, 91?92, 99?111, 102?104, 101?102, 104, 100?102, 99, 103?104 museum collaboration, 80?84 Naskapi people, 64?65 Nenet people, 70 observations of climate change, 129?139 repatriation efforts for, 71, 80, 84?86, 121, 123?125 research contributions, 79?88 selling or bartering, 94, 99?100, 101 Needell, Allan, 26 Nelson, Edward W., 64?65, 68?71, 80, 100 Neumayer, Georg von, 5 Newcomb, Simon, 17?18 32_Index_pg395-405_Poles.indd 40132_Index_pg395-405_Poles.indd 401 11/17/08 10:33:05 AM11/17/08 10:33:05 AM 402 INDEX NMAI. See National Museum of the American Indian NMNH. See National Museum of Natural History Nobel, Albert, 51 Nonnative species. See Biological invasion Nonpelagic development, 181?192 ACC hypothesis, 185?187, 192 adaptation, 182?183, 192 enhanced speciation, 185?187, 192 extinction, selective, 184?185, 191 Thorson?s rule, 181?182, 187 Northwest Passage, 353 NSF. See National Science Foundation Octopods, abundance, 197?202 depth range, 198?200 two-sample t-tests of, 200 Odishaw, Hugh, 28, 31 Oldmixon, George Scott, 93 Olofsson, Johan, 53 Olsen, Ed, 388?389 Once Round the Sun (Fraser), 28 Oozeva, Conrad, 137 Operation Highjump, 59, 241 Orton, William, 17 Pack ice, 135?136, 137, 138, 267. See also Krill, Antarctic CDOM source, 330 diving, 243, 244, 245, 248?249, 290?292 emperor penguins, 268 seals, 336, 340 Palmer Long-Term Ecological Research (LTER), ix, xiv, 289, 291?295 Papike, Jim, 389 PAR. See Photosynthetically active radiation Pelagic calanoid copepods, 143?169 abundance, 146, 148 bathypelagic, 156, 168?169 bipolar distribution, 159?160 bipolar species pairs, 166?169 classifi cation, 146 common, 160?162 deepwater, 146, 147, 148, 151?153, 153?159 endemicity, 146, 162?164 epipelagic, 146, 148, 150?151 evolution, 164?169 exoskeleton, 147?148, 169 expeditions to collect, 144 geographic distribution, 147 inshore, 146, 148, 149?150 mesopelagic, 146, 168?169 methods of collecting and studying, 145?148, 169 morphology, 147?148, 164?166, 169 Southern Ocean, 144, 148?149 sibling species pairs, 164?165, 167?168 taxonomy, 144?145, 173?179 Pelagic communities, link with benthic, 218?219 Pelagic development nonpelagic vs., 181?192 selective extinction, 184?185, 191 Personne, Mark, 52 Peters, C. H. F., 16?17 Phaeocystis antarctica, 300, 303?304, 306, 319?331. See also Chromophoric dissolved organic matter Phillip, John, Sr., 82 Photosynthetically active radiation (PAR). See also Kelp, arctic CDOM, 319?320, 322?323 krill population dynamics, 293?295 permanent-site studies, 274?275, 278?280 phytoplankton and bacteria, 299?306 Southern Ocean, 310 Phytoplankton. See also Photosynthetically active radiation CDOM source, 330 Ross Sea, 299?306 Pitul?ko, Vladimir, 70 Planet Earth (fi lm), 29 Plateau Observatory (PLATO), 377?378, 384?385 Point Barrow, Alaska. See Barrow, Alaska Polar front theory, 7?8 Polar Record (journal), 8 Polar research. See also Arctic studies; Climate studies; Cooperation, scientifi c; Diving; International Polar Year; Meteorites, Antarctica Polynyas. See Chromophoric dissolved organic matter; Ross Sea Pomerantz, Martin A., 360?361 Powell, John Wesley, 65?66, 118 Powell, Joseph S., 93 PreHEAT, 378?379, 385 Prydz Bay/East Antarctic shelf, 312 Radford, S., 362 Rainey, Froelich, 104 Ray, Patrick Henry, 19, 92, 93, 94, 96, 99?100 Rearden, Alice, 84, 86 Rearden, Noah, 86 Repatriation. See also National Museum of Natural History human remains, 71, 121 knowledge, 71, 123?125 visual, 80, 84?86 Richter, Henry L., 45 Rideout, E. B., 7 Ripley, S. Dillon, 68 Rivers, Neva, 84 Robbins, Rob, 242 Rockets, 8?9, 25, 37?46 32_Index_pg395-405_Poles.indd 40232_Index_pg395-405_Poles.indd 402 11/17/08 10:33:05 AM11/17/08 10:33:05 AM INDEX 403 Rogick, Mary, 206 Ronne, Finn, 55 Ronne, Jackie, 55 Roosevelt, Franklin, 54 Rose, Donna, 83 Ross, James Clark, 15 Rossby, Carl Gustav, 26 Rosse, Irving, 64 Ross Ice Shelf, 8, 151 Ross Sea, 206, 207, 211, 218, 299?306, 319?331, 336. See also Chromophoric dissolved organic matter; Phaeocystis antarctica; Phytoplankton seals, 266?267 emperor penguins, 268 productivity, 312?316 Russell, Sara, 390 Russia, 70. See also Soviet Union Russian Academy of Sciences, 63 Russian Institute of Cultural and Natural Heritage, 133?134 R/V Challenger (1873?1876), 144, 181, 187, 189 R/V Hero (1968?1982), 206 R/V Nathaniel B. Palmer, 302, 321 R/V Polarstern, 197?202, 290 R/V Proteus, 272 Sabine, Edward, 15, 16, 17 Saclamana, Marie, 100 Salinity kelp productivity, 272, 274, 278, 281, 282 nonpelagic development, 184 Satellites cosmic microwave background (CMB), 360 IGY, 9, 23, 24, 36, 37 NASM collection, 38?42 Nimbus 7, 310 oceanographic studies, 309?310, 320 ORBView-2, 310 TV-3, 39, 40 Schoolcraft, Henry, 62 Science?The Endless Frontier (Bush), 25?26 Sco-Cen star-forming region, 384 Scotia Arc region, 186?187, 192, 206 Scott, Robert Falcon, 51 Scott Polar Research Institute, 18 Scripps Institution of Oceanography, 241?242 Scuba diving. See Diving Sea Ice Knowledge and Use: Assessing Arctic Environmental and Social Change (SIKU), 133?138 Sea-ice microbial communities (SIMCOs), 288?289, 290?291, 292, 293, 294, 295 Sea ice. See also Climate change; Emperor penguins; Weddell seals; Yupik Eskimos formation, 66?67 Savoonga, Alaska, observations from, 134?135, 138 Sea level, rise of, 11 Seals. See also Weddell seals climate change, 138 diving behavior, 267?268 diving hazard, 249 diving physiology, 268?269 krill in diet, 286 lactation, 336?337 Yup?ik language, 86 Sea-Viewing Wide Field-of-View Sensor (SeaWiFS), 310?311, 328 Senungetuk, Ronald, 69 Serov, Sergei, 70 Shumaker, Brian, 59 SIKU. See Sea Ice Knowledge and Use: Assessing Arctic Environmental and Social Change Silook, Paul, 132, 134 Simmonds, Doreen, 100 Simpson, John, 91?92, 94, 108 Simpson, Thomas, 90 Singer, S. Fred, 38 Siple, Paul, 24, 54 Sky noise, 362 Smith, Middleton, 92, 93 Smithsonian Institution. See also Arctic Studies Center; Exhibits; Expeditions arctic studies, 61?74 Alaska connection of, revitalizing, 70?72 Eskimo art, public discovery, 67?70 ethnological collecting, 62?65, 89?96 historical contributions, 61?62 IPY-1 participation, 61?62, 64?65, 89?96 IPY-4 participation, 72?74 meteorite program participation, 389?393 professional staff, development of, 65?66 purchasing or bartering by, 94, 99?100, 101 Scientifi c Diving Program, xiv, 242 Smithsonian Astrophysical Observatory (SAO), xiv Smithsonian Environment Research Center (SERC), xiv Smithsonian Oceanographic Sorting Center (SOSC), xiv, 144, 206 whaling collections, 99?111 Yup?ik contributions to research, 79?88 Solar radiation. See Chromophoric dissolved organic matter; Ross Sea Sonnenfi eld, Joseph, 95?96 S?rlin, Sverker, 51 Southern Ocean. 143?169, 181?192, 309?317. See also Pelagic calanoid copepods; Ross Sea British Discovery Committee, 144 brooding species, 181?192 32_Index_pg395-405_Poles.indd 40332_Index_pg395-405_Poles.indd 403 11/17/08 10:33:07 AM11/17/08 10:33:07 AM 404 INDEX crustaceans, 191?192 iron concentration, 314?315 krill as foundation species, 285?287 productivity, 309?317 satellite studies, 309?310 temperature, 310?311, 313?314, 315?317 South Pole, as observatory site, 361?364 South Shetland Islands bryozoan study, 207?219 octopod abundance, 197?202 Soviet Union cultural exchange, 69?70 IGY participation, 29?31, 36 Soviet Antarctic expeditions (1955?1958), 144 Space age origins, and IGY, 35?47 Special Sensor Microwave Imager (SSM/I), 311 Spectroscopic imaging, high-resolution, 375 Spencer, Robert F., 95 Splettstoesser, John, 59 Spude, Catherine, 55 Spude, Robert, 55 Sputnik 1, 37, 38?39 SS Vikingen expedition (1929?1930), 144 Standards of the Conduct of Scientifi c Diving, 242 Star birth (formation), 381?386 Chamaeleon region, 382 HEAT observations, 374?375 Lupus region, 382?384 observing, from Antarctic Plateau, 381?386 rho Ophiuchi region, 382, 384 Star formation. See Star birth Stefansson Sound Boulder Patch, kelp productivity, 271?283 STELLA ANTARCTICA, 384, 385 Stevens, Ann, 69 Stevens, Ted, 69 Stewart, James R., 241?242 Stewart, T. Dale, 67, 69 Strindberg, Nils, 51, 53 Sturtevant, William C., 68 Submillimeter Polarimeter for Antarctic Remote Observing (SPARO), 363 Subtropical Convergence, 145, 147 Sullivan, Walter, 31, 35 Sunyaev?Zel?dovich effect (SZE), 363 ?Superstitions of the Eskimo? (Smith), 93 Swan, James G., 64 Tasmanian Gateway, 186?187 Taylor, C. J., 14, 15 Telegraph astronomy, 16?17 Meteorological Project, 3, 15?16 polar research, 18, 63 Telescopes. See also Antarctic Submillimeter Telescope and Remote Observatory; High Elevation Antarctic Terahertz Telescope Antarctic Searching for Transiting Extrasolar Planets (ASTEP), 385 Center for Astrophysical Research, x, 361 Degree-Angular Scale Interferometer (DASI), 361, 363 International Robotic Antarctic Infrared Telescope (IRAIT), 385 Project Moonwatch, 38 Python, 361, 362 QUEST (Q and U Extra-Galactic Sub-mm Telescope), 364 South Pole, x, xiv, 364 South Pole Infrared Explorer (SPIREX), 361 Viper, 361, 363 White Dish, 361 Temperature. See also Cold adaptation biological invasion, 350?355 cosmological observations, 360 diving, 250 kelp productivity, 272, 274, 278, 280?281, 282 nonpelagic development, 183 Southern Ocean, 310?311, 314 Terra Nova Bay, 266?267 Terra Nova collections, 144 Thomas, Twyla, 390 Thompson, Tommy, 241 Thorson?s rule, 181?182, 187 Toovak, Kenneth, 100, 106?107, 108, 110 Total suspended solids (TSS). See Kelp, arctic Tousey, Richard, 45?46 Traditional ecological knowledge (TEK), 133 Tupek, Karen, 55 Turner, J. Henry, 100 Turner, Lucien collections, 80 ethnological studies, 64, 65, 71, 117?121, 131?132 Fort Chimo (Kuujjuaq), 119?120 and Innu/Inuit peoples, 65, 119?121 Turner, Mort, 389 ?Tusking,? 238 Udvar Hazy Center, 46 Ultraviolet light CDOM screening, 319?320 productivity, 299?306 UNESCO, 28, 31 Ungava Bay, 64, 117?118 U.S. Antarctic Diving Program, x, 241?251 U.S. Antarctic Meteorite Program, ix, xiv, 388?394 U.S. Antarctic Research Program (USARP) bryozoans, 206?208, 218 pelagic calanoid copepods, 144?145, 146, 169 32_Index_pg395-405_Poles.indd 40432_Index_pg395-405_Poles.indd 404 11/17/08 10:33:08 AM11/17/08 10:33:08 AM INDEX 405 U.S. Naval Support Force Antarctica, 241 U.S. Research and Development Board, 25?26, 29 USNS Eltanin collections, 144?145, 148, 206 Van Allen, James, 24, 35, 44?45 VanStone, James, 70 Venus, transits of, 14, 17?18, 19 American Transit of Venus Commission, 17?18 weather, 17?18 Voznesenskii, Ilya G., 63 Warming. See Climate change Waterman, Alan, 28 Waugh, Leuman M., 83 Weather studies. See Climate studies Weaver, Robert, 55 Weber, Wilhelm, 14 Weddell seals (Leptonychotes weddellii), x, xiv, 265?270, 335?343 Welzenbach, Linda, 390 Western Union Telegraph Company, 17, 18, 63 Weyprecht, Karl, 5, 11, 13 Whales. See also Narwhals; Whaling beluga whales, 102, 138 bowhead whales, 99, 100, 102, 138 climate change, 135?138 diving hazard, 249 gray whales, 138 krill in diet, 286 lactation, 336?337 Whaling annual cycle, 102?104 ceremonies and celebrations, 101?102, 104 climate change, 135?138 contact and change, 100?102 contemporary, 99 elders? interpretation of collection, 99?100, 104?110 fi rearms use, 95, 101 I?upiaq society, 99?111 I?upiaq villages, 99 IWC quota, 102 kayak use, 83?84 modern methods, 103?104 preparations, 102?103 specifi c object discussions, 104?110 and women, 102?103, 108?109 Whymper, Frederick, 63 Wild, Heinrich, 5, 11 Williams, George, 122 Wilson, A. C., 256 World Climate Research Programme (WCRP), 10 World Meteorological Organization (WMO), 26?27, 29 World War I, 6?8 World War II, 8?9, 54 Yanai, Keizo, 388 Yupik Eskimos, 129?139 Yup?ik Eskimos. See also Native peoples art, public discovery of, 67?70 elders, museum collaboration with, 80?84 ethnological collecting, 65?66 homeland, 80, 81 language, 85, 86?87, 135 objects in collections, 79?80, 83?85, 86 pride and self-respect, 82 research contributions, 79?88 tools and technology, 85?87 visual repatriation, 80, 84?86 Zeitlin, Sam, 44 32_Index_pg395-405_Poles.indd 40532_Index_pg395-405_Poles.indd 405 11/17/08 10:33:08 AM11/17/08 10:33:08 AM