Biology of the Southern Ocean

622 Pages·2007·20.29 MB·English

by  George A. Knox

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.
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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,
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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.
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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.
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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.
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