30° 0° 30° N T A R C TIC—CONVERGENCE A Prince Edward Bouvetaya Marion Island Island South South Sandwich Georgia Islands 60° lies 60° Crozet Falkland South Orkney Sanae Neumayer SOUTH Islands Islands AMERICA Signy Syowa South QUEEN MAUD Shetland 75°S LAND lies Islands Larsen Halley ENDERBY McDonald Is Kerguelen Ice Shelf LAND Mawson ANTA RoIcnene Prydz Bay HIselaanrdd RT Shelf GREATER IC ANTARCTICA Davis 90° Siple PENIN LESSER Amundsen Mirny 90° Peter I Island SULA ANTARCTICA Scott Oasis WILKES MARLIAEN BDYRD Ross LAND Casey Ice Shelf VICTORIA ROSS LAND SEA McMurdo Scott 75°W 70° 65° 60° 55°W Bellingshausen Hallett Frei Dumont Arctowski Balleny D'Urville Deception I Islands Scott Island Palmer 65°S Faraday 60°S 120° Rothera Almirante Brown Macquarie Island Campbell Island Auckland Island 70°S Fossil 45°S Bluff NEW AUSTRALIA 80°W 50°W 180° ZEALAND 150° Antarctica is both a continent and an ocean. The continent is inextricably linked to the vast extent of the Southern Ocean surrounding it, an ocean that supports a teaming array of life, ranging from microscopic plants that form part of the phytoplanktonresponsibleforprimaryproductiontothemightywhales,ourlargestmarinemammals.Whilstourknowledge of the biology of these waters has been accumulating since the first voyages of discovery in the 18th century, there is yet to be a comprehensive, single-author synthesis of the current state of that knowledge. This book sets out to correct that deficiency. q2006byTaylor&FrancisGroup,LLC q2006byTaylor&FrancisGroup,LLC q2006byTaylor&FrancisGroup,LLC q2006byTaylor&FrancisGroup,LLC Preface Our knowledge of the biology of the seas surrounding the continent. The voluminous reports of these Discovery Antarctic continent began with descriptions and specimens expeditionslaidtheindispensablefoundationforsubsequent broughtbackbyearlynaturalistsonvoyagesofdiscoveryin studies. theSouthernOcean. Prominentamongthese werethe early Southern Ocean marine research entered a new phase FrenchnavigatorssuchasBouvetdeLozier,whodiscovered followingtheestablishment,in1957,oftheSpecial(laterthe Bouvet Island in 1739, and Yves Joseph de Kerguelen- Scientific) Committee on Antarctic Research (SCAR). This Tremarez, who discovered the island that bears his name initiatedthemoderneraofSouthernOceanresearchinwhich some30yearslater.Thesewerefollowedbythevoyagesof many countries played roles, notably the Soviet Antarctic Captain Cook, who, in 1772, reached 718 100 S in the ExpeditionsintheVitezandObandtheUnitedStatesEltanin Bellingshausen Sea. In 1819, the Russian explorer Admiral cruises.OthercountriessuchasGreatBritain,France,South Bellingshausenwasfirsttotowanetatthesternofhisship Africa,Chile,Argentina,AustraliaandJapanwerealsovery and he found that it collected more organisms during the activeduringthisperiod.In1972,acoordinatedinternational nightthanduringtheday.Hewasprobablythefirsttocollect researchprogramwasinitiatedwiththeestablishmentofthe theAntarcticakrill,Euphausiasuperba. BIOMASS (Biological Investigations of Antarctic Systems The first serious scientific marine research was carried andStocks)Program(El-Sayed1996). out by James Eights who sailed in 1929 with the American My own involvement in Antarctic research began with sealingCaptainsPalmerandPendleton.Eightsdescribedthe a visit to McMurdo Sound in 1960 where one of my staff naturalhistoryoftheSouthShetlandIslands.Hisbestknown in the Department of Zoology at the University of discoverywasofaten-leggedpycnogonid.Soonafter,there Canterbury, Dr. Bernard Stonehouse, was initiating a followed a series of national expeditions led by Dumont research program on Adelie penguins and McCormick D’Urville of France (1837–1840), Charles Wilkie of the skuas. This led to the establishment of the University of United States (1838–1842), and Sir James Clark Ross of Canterbury Antarctic Research Unit (Knox 1986, 1988) Britain (1939–1943). Among the naturalists who accom- that carried out continuous summer research in the panied these expeditions, the most notable were J.D. Dana, McMurdo Sound region over the period 1960–1983, the on the Wilkie expedition, and J.D. Hooker, on the Ross last twelve years of which I was director of the unit. expedition.Theseexpeditionsallmadeextensivecollections During the summers of 1969 and 1970, I initiated an ofmarineorganisms. inshore marine research program that continued until the TherewasarenewalofAntarcticexplorationattheend 1982–1983 season. Throughout the operation of the unit, of the nineteenth century. Major expeditions included the some sixty research students participated in its activities. British (1899–1900), the Belgian (1898), the Swedish Interaction with these enthusiastic young minds had a National Expedition (1902–1904), the Scottish National significant influence on many of the concepts developed in Antarctic Expedition (1902–1904), the German expedition The Biology of the Southern Ocean. I also have had the (1901–1903)intheGauss,andtheFrenchExpeditioninthe good fortune to become involved in international Antarctic PourquoiPas(1908–1910).Alloftheseexpeditionsresulted science activities, first as a member and secretary of the intaxonomicstudiesthatlaidthefoundationforlaterstudies SCAR Biology Working Group on Biology and as a ofthemarinefloraandfauna. member of SCAR since 1969. I was a member of the EcologicalstudiesintheSouthernOceanbeganwiththe Group of Specialists on the Living Resources of the work of naturalists attached to land-based, over-wintering Southern Ocean from its inception and I attended the first parties such as those of Sir Ernest Shackleton’s 1907–1909 two meetings of CCAMLR (Convention on the Conserva- expedition and Captain Scott’s two polar expeditions. tion of Antarctic Marine Living Resources). These Antarctic marine studies received a tremendous impetus activities brought me into contact with a wide range of fromtheinvestigationsthatbeganin1925withthestudyof Antarctic scientists from all the SCAR countries. whalecarcassesatthewhalingstationsatGrytvikeninSouth The suggestion for The Biology of the Southern Ocean Georgia.Theworkofthescientistsexpandedtoinclude,not wasmadebyDr.BernardStonehouseand,withouthisencour- onlystudiesonwhaledemographyandecology,butalsoof agement, it would not have been written. Discussions with the physical, chemical and biological oceanography of the AntarcticcolleaguesthroughouttheSCARcommunityhave Southern Ocean in order to gain an understanding of the hadaprofoundinfluenceonitsdevelopment.AmongsttheseI factorsinfluencingthedistribution,reproduction,andgrowth particularlywishtothankProfessorSayedEl-Sayed,thelate ofwhalestocks.Theseinvestigationscontributedtothefirst Sir George Deacon, Professor A.L. DeVries, Professor M. long-term studies of the Antarctic pelagic ecosystem, and Fukuchi,SirMartinHoldgate,ProfessorT.Hoshiai.Dr.J.C. they were the first that extended right around the Antarctic Hureau,Dr.K.B.Kerry,Dr.Y.Naito,thelateDr.T.Nemoto, q2006byTaylor&FrancisGroup,LLC Dr. J. Oliver, Dr. J. Warham, and Professor E.C. Young. publicationofthefirsteditionofTheBiologyoftheSouthern Accesstoliteraturewasgreatlyfacilitatedbyperiodsspentin Ocean.Whilesomeoftheinformationinthefirsteditionis thelibrariesoftheBritishAntarcticSurveyandtheNational retained, it is essentially a new account. The task of Institute of Polar Research, Tokyo. Dr. D.W.H. Walton summarizing the large volume of research has proved a reviewedallchaptersoftheoriginaleditioninvariousdrafts difficult task. Most of the figures in the text have been and I am grateful for his assistance and encouragement. I redrawnfromtheoriginals. would also like to thank Dr. Maria Murphy and the staff of Thebookcommenceswithadescriptionofthephysico- Cambridge University Press for their patience and support chemical environment of the Southern Ocean and then duringthepreparationofthefirsteditionandforseeingthe follows a logical sequence covering phytoplankton and projectthroughtocompletion. primaryproduction,theseaicemicrobialcommunities,and The first edition of The Biology of the Southern Ocean the secondary consumers, the zooplankton. There is an waspublishedin1993andwaswellreceived.Theprintrun extended chapter on the biology and ecology of Antarctic was sold out a number of years ago. Some 13 years have krill, in view of its central position in the Southern Ocean passedsincethenandaconsiderableamountofresearchhas foodweb.Additionally,krillhasbeenthesubjectofintensive been carried out during this period. The original SCAR research programs over the past decades, especially during countriesengagedinmarineresearchintheSouthernOcean theBIOMASSProgrammesandsubsequentlybyCCAMLR. have been joined by other countries including China, A series of chapters consider the higher consumers, nekton Ukraine, Uruguay, Korea, Ecuador, India, Peru and Spain, (with an emphasis on cephalopods), fish, seals, whales and resultinginamuchincreasedresearcheffort. seabirds. A series of chapters then follow on selected The last decade in particular had seen the initiation of ecosystem components: the benthic communities, life several scientific programs to study phenomena and beneath the fast ice and ice shelves, recent advances in processes of global significance in which the Southern understanding decomposition processes, and the role of Ocean plays a key role. Some of the major programs are bacteriaandprotozoa.Thesearefollowedbyanattemptata listedbelowandbriefdescriptionsoftheobjectivesofthese synthesis of ecosystem dynamics, with an emphasis of the aregiveninEl-Sayed(1991). pelagic ecosystem and then three chapters dealing with - AntarcticMarineLivingResources(AMLR)program resourceexploitation,theimpactofsuchexploitationonthe (1986-present) marine ecosystem, and the problems involved in the - AntarcticMarineEcosystemResearchattheIce-Edge management of the living resources. Three new chapters Zone(AMERIEZ)(1983–1988) havebeenaddedtothesecondeditionexploringtheimpact - Research on Antarctic Coastal Ecosystem Rates of increased UV radiation, human impact on the marine Program(RACER)(1986–1993) environment,andtheimpactofglobalwarmingonSouthern - Long-Term Ecological Research (Palme LTER) Oceanmarineecosystems. (1980-present) As can be seen by the reference list at the end of this - Southern Ocean Joint-Global Ocean Ecosystem volume, there is a considerable body of recent literature on Dynamics(SO-GLOBEC)(1991–2009) thebiologyoftheSouthernOcean.Becauseofthegrowthof - Coastal and Shelf Ecology of the Antarctic Sea-Ice publishedresearch,Ihavehadtobeselectiveinthematerial Zone(CS-EASIZ)(1993–2004) thatisincluded.Exampleshavebeencarefullychosenfrom - AntarcticPack-IceSeals(APIS)(1995-present) the pool of published research to illustrate the concepts - EuropeanPolarsternStudy(EPOS)(1988–89) discussed.Thereare,doubtless,othersthatcouldhavebeen - Research on Ocean-Atmosphere Variability and used and I apologize to authors whose work has not been Ecosystem Response in the Ross Sea (ROAVERRS) included. (1996–1998) In the preparation of this volume I have been supplied Themulti-disciplinary,multi-nationalcruisesoftheR/V with advanced copies of papers prepared or accepted for Polarstern have been of particular importance in recent publicationandIamgratefultoallwhosuppliedthem.Iam years, especially the ANDEP cruises (2002–2005) that alsoappreciativeofthesupportandencouragementofJohn investigated the benthic fauna of the deep Weddell Sea. Sulzycki and the staff of CRC Press, in particular Pat Over the period 2000–2002, a series of iron fertilization Roberson, during the preparation of this edition and for experiments (SOIRE, SOFex and EISENEX) investigated seeingtheprojectthroughtothecompletionofahighquality thepotentialroleofironincontrollingalgalproductioninthe product. Southern Ocean. Recently, the New Zealand National This book is an attempt to synthesize the available Institute of Water and Atmospheric Research (NIWA) information into a coherent account of one of the most investigated the richness and diversity of the benthos in a fascinating systems on the globe. I hope that it will prove latitudinal gradient in the western Ross Sea in relation to useful to advanced undergraduates and to professionals broader-scaleenvironmentalfactorsandlocal-scalevariation engaged in Antarctic marine research, as well as to all inproductivitysources. interestedorinvolvedinAntarcticmarineconservationand Astheresultoftheresearchactivitieslistedabove,there management. It will have been worthwhile if it stimulates hasbeenaconsiderablevolumeofpublishedresearchonthe otherstoworkonthefascinatingaspectsofAntarcticmarine biology of the Southern Ocean in the 13 years since the researchthathavebeendiscussedinthisbook. q2006byTaylor&FrancisGroup,LLC Acknowledgments IwouldfirstlyexpressmyindebtednesstothemanyinternationalAntarcticscientists,someofwhomhavebeenlistedinthe Preface, who Ihave interacted withover the years.Discussions with themhave influenced the developmentof many of the conceptsdicussedinthisbook.IacknowledgetheoutstandingcontributionofCambridgeUniversityPress,inthepreparation ofthefirsteditionofthisbook.Ioweagreatdebttothestaffmembersandstudentswhoparticipatedintheresearchactivitiesof theUniversityofCanterburyAntarcticResearchUnit. IoweagreatdealtoJohnSulzyckiandthestaffofCRCPress,inparticularPatRobersonfortheirpatienceandsupport duringthepreparationofthissecondedition.TogetherwiththestaffofAldenPressServiceswhotypesetthisbooktheyhave producedahighqualityproduct. q2006byTaylor&FrancisGroup,LLC q2006byTaylor&FrancisGroup,LLC Author GeorgeA.Knox,C.N.Z.M.,M.B.E.,F.R.S.N.Z.,washead University of Canterbury Antarctic Research Unit. He of the Department of Zoology, University of Canterbury, has participated in many field expeditions, including the Christchurch, New Zealand, from 1959 to 1976. He is now ChathamIslands1954Expedition(leader);theRoyalSociety professoremeritusofzoology. of London Darwin Centennial Expedition to southern Chile Professor Knox was born in New Zealand and received (marine biologist and deputy leader); thirteen summer hiseducationattheUniversityofCanterbury,wherehewas expeditionstoMcMurdoSound,Antarctica;theestablishment appointed a staff member in 1948. He has been a visiting of the Snares Islands Research Program (he participated fellow at the East-West Center, Honolulu, and a visiting in three field expeditions); and participation in field professor at the Department of Oceanography, Texas A&M expeditions to Campbell and Auckland Islands. He has University and the Department of Environmental Engin- published over 100 scientific papers and 28 environmental eering,UniversityofFlorida,Gainesville.Hehasvisitedand reports,writtenfivebooks,andeditedandco-authoredthree worked in laboratories in the U.S., Canada, Chile, Japan, othervolumes. Australia,WesternEurope,theUSSR,andChina. Professor Knox has received a number of awards and Professor Knox’s research has been wide ranging and fellowshipsforhiscontributionstoscience,includingFellow includes: (1) the systematics and distribution of polychaeta oftheRoyalSocietyofNewZealand(FRSNZ),1963;Hutton with special reference to New Zealand and Antarctica; (2) Medal, Royal Society of New Zealand, 1978; Conservation rocky shore intertidal ecology and biogeography; (3) the Trophy,NewZealandAntarcticSociety;HonoraryMemberof ecologyandconservationofislands;(4)studiesonthepelagic ScientificCommitteeonAntarcticResearch,1982;memberof and benthic ecosystems beneath the sea ice in McMurdo theMostExcellentOrderoftheBritishEmpire(MBE),1985; Sound,Antarctica;and(5)estuarineandcoastalecologyand theNewZealandAssociationofScientists’SirErnestMarsden management. He established and directed the Estuarine Medal for Service to Science, 1985; and Companion of the Research Unit in the Department of Zoology and the NewZealandOrderofMerit(CNZM),2001. q2006byTaylor&FrancisGroup,LLC q2006byTaylor&FrancisGroup,LLC