ANTARCTIC American GeophysicaUl nion RESEARCH SERIES Foundations for Ecological Research West of the Antarctic Peninsula Antarctic Research Series Volumes 1 Biology of the Antarctic Seas I Milton O. Lee 29 Upper Atmosphere Research in Antarctica (Ed.) L. J. Lanzerotti, C. G. Park (Eds.) 2 Antarctic Snow and Ice Studies M. Mellor 30 Terrestrial Biology II B. Parker (Ed.) 3 Polychaeta Errantia of Antarctica O. Hartman 31 Biology of the Antarctic Seas IX L.S. (Ea.) Kornicker (Ed.) 4 Geomagnetisma nd Aeronomy A.H. Waynick 32 Biology of the Antarctic Seas X L.S. (Ea.) Kornicker (Ed.) 5 Biology of the Antarctic Seas II G.A. Llano 33 Dry Valley Drilling Project L.D. McGinnis (Ea.) (Ed.) 6 Geology and Paleontology of the Antarctic 34 Biology of the Antarctic Seas XI L.S. J. B. Hadley (Ed.) Kornicker (Ed.) 7 Polychaeta Myzostomidae and Sedentaria of 35 Biology of the Antarctic Seas XII D. Pawson Antarctica O. Hartman (Ed.) (Ed.) 8 Antarctic Soils and Soil Forming Processes 36 Geology of the Central Transantarctic J. C. F. Tedrow (Ed.) MountainsM . D. TurnerJ,i F. Splettstoesser 9 Studies in Antarctic Meteorology M.J. Rubin (E&.) (Ea.) 37 Terrestrial Biology III B. Parker (Ed.) 10 Entomology of Antarctica J.L. Gressit (Ed.) 38 Biology of the Antarctic Seas XIII [crinoids, 11 Biology of the Antarctic Seas III G.A. Llano, hydrozoa, copepods,a mphipoda] L.S. W. L. Schmitt (Eds.) Kornicker (Ed.) 12 Antarctic Bird Studies O. L. Austin, Jr (Ed.) 39 Biology of the Antarctic Seas XIV L.S. 13 Antarctic Ascidiacea P. Kott (Ed.) Kornicker (Ed.) 14 Antarctic Cirripedia W. A. Newman, A. Ross 40 Biology of the Antarctic Seas XV L.S. (mas.) Kornicker (Ed.) 15 Antarctic OceanologyI L. Reid (Ed.) 41 Biology of the Antarctic Seas XVI L.S. 16 Antarctic Snow and Ice Studies II A.P. Crary Kornicker (Ed.) (Ea.) 42 The Ross Ice Shelf.' Glaciology and Geophysics 17 Biology of the Antarctic Seas IV G.A. Llano, C. R. Bentley,D . E. Hayes (Eds.) I. E. Wallen (Eds.) 43 Oceanology of the Antarctic Continental Shelf 18 Antarctic Pinnipedia W. H. Burt (Ed.) S. Jacobs( Ed.) 19 Antarctic Oceanology II: The Australian-New 44 Biology of the Antarctic Seas XVII [benthic Zealand Sector D. E. Hayes (Ed.) satiation, brittle star feeding, pelagic shrimps, 20 Antarctic Terrestrial Biology G.A. Llano marine birds] L. S. Kornicker, (Ed.) (Ea.) 45 Biology of the Antarctic Seas XVIII, Crustacea 21 Recent Antarctic and Subantarctic Brachiopods Tanaidacea of the Antarctic and the M. W. Foster (Ed.) Subantarctic 1. On Material Collected at 22 Human Adaptability to Antarctic Conditions Tierra del Fuego, Isla de los Estados, and the E. K. Eric Gunderson (Ed.) West Coast of the Antarctic Peninsula L.S. 23 Biology of the Antarctic Seas V D.L. Pawson Kornicker (Ed.) (Ea.) 46 Geological Investigations in Northern Victoria 24 Birds of the Antarctic and Sub-Antarctic Land E. Stump (Ed.) G. E. Watson 47 Biology of the Antarctic Seas XIX [copepods, 25 Meterological Studies at Plateau Station, teleosts] L. S. Kornicker (Ed.) Antarctica J. Businger( Ed.) 48 Volcanoes of the Antarctic Plate and Southern 26 Biology of the Antarctic Seas VI D.L. Pawson Oceans' W. E. LeMasurierJ, . W. Thomson (ma.) (Eds.) 27 Biology of the Antarctic Seas VII D.L. 49 Biology of the Antarctic Seas XX, Antarctic Pawson (Ed.) Siphonophores From Plankton Samples of the 28 Biology of the Antarctic Seas VIII D.L. United States Antarctic Research Program Pawson, L. S. Kornicker (Eds.) L. S. Kornicker (Ed.) 50 Contributions to Antarctic Research I D. /-/. 61 Antarctic Meteorology and Climatology: Studies Elliot (Ed.) Based on Automatic Weather Stations D. /-/. 51 Mineral Resources Potential of Antarctica Bromwich, C. R. Stearns (Eds.) J. F. SplettstoesseGr., A.M. Dreschhoff( Eds.) 62 Ultraviolet Radiation in Antarctica: 52 Biology of the Antarctic Seas XXI [annelids, Measurements and Biological Effects C.S. mites, leeches] L. S. Kornicker (Ed.) Weiler,P . A. Penhale (Eds.) 53 Contributions to Antarctic Research II D.H. 63 Biology of the Antarctic Seas XXIII, Antarctic Elliot (Ed.) and Subantarctic Pycnogonida: Ammotheidae 54 Marine Geological and GeophysicalA tlas of the and Austrodecidae S. D. Cairns (Ed.) Circum-Antarctic to 30ø S D. E. Hayes (Ed..) 64 Atmospheric Halos W. Tape 55 Molluscan Systematics and Biostratigraphy 65 Fossil Scleractinian Corals From James Ross Lower Tertiary La Meseta Formation, Seymour Basin, Antarctica H. F. Filkorn Island, Antarctic Peninsula J. D. Stilwell, W. J. 66 Volcanological and Environmental Studies of Zinsmeister Mt. Erebus P. R. Kyle (Ed.) 56 The Antarctic Paleoenvironment: A Perspective 67 Contributions to Antarctic Research IV D.H. on Global Change, Part One J.P. Kennett, Elliot, G. L. Blaisdell (Eds.) D. A. Warnke (Eds.) 68 Geology and Seismic Stratigraphy of the 57 Contributions to Antarctic Research Ill D.H. Antarctic Margin A. K. Cooper,P . F. Barker, Elliot (Ed.) G. Brancolini (Eds.) 58 Biology of the Antarctic Seas XXII S.D. 69 Biology of the Antarctic Seas XXIV, Antarctic Cairns (Ed.) and Subantarctic Pycnogonida: Nymphonidae, 59 Physical and BiogeochemicalP rocessesi n Colossendeidae, Rhynchothoraxida, Antarctic Lakes W.J. Green, E. I. Friedmann Pycnogonidae,P hoxichilidiidae, Endeididae, (Eds.) and Callipallenidae S. D. Cairns (Ed.) 6O The Antarctic Paleoenvironment: A Perspective 7O Foundations for Ecological Research West of on Global Change, Part Two J.P. Kennett, the Antarctic Peninsula R. M. Ross, E. E. D. A. Warnke (Eds.) Hofmann, L. B. Quetin (Eds.) ANTARCTIC Volume 70 RESEARCH SERIES Foundationsf or EcologicalR esearch West of the Antarctic Peninsula Robin M. Ross Eileen E. Hofmann Langdon B. Quetin Editors (cid:127) AmericGaeno physUicnaiol n Washington, D.C. 1996 ANTARCTIC Volume 70 RESEARCH SERIES FOUNDATIONS FOR ECOLOGICAL RESEARCH WEST OF THE ANTARCTIC PENINSULA Robin M. Ross, Eileen E. Hofmann, and LangdonB . Quetin,E ditors Publishedu ndert he aegiso f the Board of Associate Editors, Antarctic ResearchS eries RodneyM . Feldmann,C hairman JohnB . Anderson,R obert A. Bindschadler, David H. Bromwich, Nelia W. Dunbar, StanleyS . JacobsJ, erryD . Kudenov,J ohnC . Priscu Library of CongressC ataloging-in-PublicationD ata Foundationsf or EcologicalR esearchW est of the Antarctic Peninsula/ Robin M. Ross,E ileen E. Hofmann,L angdonB . Quetin,e ditors. p. cm.- - (Antarctirce searcshe riesv; . 70) Includesb ibliographicarl eferences. ISBN 0-87590-891-8 (alk. Paper) 1. Ecology--Research(cid:127)Antarctica(cid:127)Antarctic PeninsulaR egion. I. Ross, Robin M. II. Hofmann, Eileen Elizabeth, 1952- IlI. Quetin, LangdonB . IV. Series. QH541.264.A6F67 1996 574.5'0998' 9---dc20 96-32835 CIP ISBN 0-87590-891-8 ISSN 0066-4634 Copyright1 996 by the AmericanG eophysicaUl nion 2000 Florida Avenue, N.W. Washington,D C 20009 Figurest,a bles,a nds horte xcerptsm ayb e reprintedin scientificb ooksa ndj oumalsi f the source is properlyc ited. Authorizationto photocopyit emsf or internaol r personaul se,o r the internalo r personaul seo f specific clients,i s grantedb y the AmericanG eophysicaUl nion for librariesa nd otheru sers registeredw ith the CopyrightC learanceC enter (CCC) TransactionaRl eportingS ervice, providedth att heb asef eeo f $01.50p erc opyp lus$ 0.50p erp agei s paidd irectlyt o CCC, 222 RosewoodD r., Danvers,M A 01923. 0066-4634/96/$01.50+0.50. This consentd oesn ot extendt o otherk indso f copying,s ucha s copyingf or creatingn ew collectivew orkso r for resale. The reproductioonf multiplec opiesa ndt he useo f full articles oar the use of extracts,i ncluding figures and tables, for commercialp urposesr equires permissionfr om the AmericanG eophysicaUl nion. Publishedb y American GeophysicalU nion With the aid of grant OPP-9414962 from the National Science Foundation Printed in the United States of America. CONTENTS The Antarctic Research Series Board of Associate Editors xi Preface Eileen E. Hofmann, Robin M. Ross,a nd Langdon B. Quetin xiii Historical Perspective Historical Perspectiveo f Researchi n the Antarctic PeninsulaR egion SayedZ . El-Sayed Habitat Description Terrestrial and FreshwaterB iotic Componentso f the Western Antarctic Peninsula RonaM I. Lewis Smith 15 Water Mass Distribution and Circulation West of the Antarctic Peninsulaa nd Including Bransfield Strait Eileen E. Hofmann, John M. Klinck, Cathy M. Lascara, and David A. Smith 61 Spatial and Temporal Variability of Western Antarctic PeninsulaS ea Ice Coverage S. E. Stammerjohna nd R. C. Smith 81 SurfaceA ir Temperature Variations in the Western Antarctic PeninsulaR egion Raymond C. Smith, Sharon E. Stammerjohn,a nd Karen S. Baker 105 Benthic Marine Habitats in Antarctica Andrew Clarke 123 Accumulationo f Glacial Marine Sedimentsi n Fjords of the Antarctic Peninsulaa nd Their Use as Late Holocene Paleoenvironmental Indicators Eugene W. Domack and CharlesE . McClennen 135 Distributions of Biological Populations Pelagica nd Sea Ice Microbial Communities David L. Garrison and SylvieM athot 155 Phytoplankton:Q uantitative and Qualitative Assessments R. R. Bidigare,J . L. Iriarte, S.-H. Kang, D. Karentz,M . E. Ondrusek,a nd G. A. Fryxell 173 Distribution of Antarctic Krill and Dominant Zooplankton West of the Antarctic Peninsula Robin M. Ross,L angdon B. Quetin, and Cathy M. Lascara 199 The Distribution of Antarctic Marine Benthic Communities Andrew Clarke 219 Midwater Fish Ecology Adolf K. Kellermann 231 Factors Controlling the Distribution of Seabirds:W inter-Summer Heterogeneityi n the Distribution of Addlie Penguin Populations William R. Fraser and WayneZ . Trivelpiece 257 The Breeding Biology and Distribution of Addlie Penguins:A daptations to Environmental Variability WayneZ . Trivelpiecea nd William R. Fraser 273 Marine Mammals of the Southern Ocean Daniel P. Costa and Daniel E. Crocker 287 Control of Biological Distributions MicrobiologicalO ceanography(cid:127)int he Region West of the Antarctic Peninsula:M icrobial Dynamics,N itrogen Cycle and Carbon Flux David M. Karl, James R. Christian, John E. Dore, and Ricardo M. Letelier 303 PhytoplanktonB iomassa nd Productivityi n the Western Antarctic PeninsulaR egion Raymond C. Smith, Heidi Dierssen,a nd Maria l/ernet 333 Factors Affecting Distribution and Abundance of Zooplankton, With an Emphasiso n Antarctic Krill, Euphausias uperba Langdon B. Quetin, Robin M. Ross, ThomasK . Frazer, and Karen L. Haberman 357 Marine Benthic Populationsi n Antarctica: Patterns and Processes Andrew Clarke 373 Human Disturbance Human Activity and Disturbance:B uilding an Antarctic Site Inventory Ron Naveen 389 Marine Disturbance-Contaminants Mahlon C. Kennicutt H and Susanne J. McDonaM 401 Marine Disturbances-CommerciaFl ishing David J. Agnew and SteveN icol 417 Summary The Western Antarctic PeninsulaR egion: Summaryo f Environmental and Ecological Processes The Palmer L TER Group 437 THE ANTARCTIC RESEARCH SERIES The AntarcticR esearchS eriesp, ublisheds ince1 963b y the AmericanG eophysicaUl nion, now comprise7s0 volumeso f authoritativoer iginarl esultso f scientifiwc ork in theh ighl atitudeos f the southernh emisphereS. eriesv olumesa re typicallyt hematicc, oncentratinogn a particulart opic or region,a ndm ay containm apsa ndl engthyp apersw ith largev olumeso f datai n tabularo r digital format. Antarctic studiesa re often interdisciplinaryo r international,a nd build upon earlier observationtso addressis sueso f naturalv ariabilitya nd globalc hange.T he standardso f scientific excellenceex pectedfo r the Seriesa re maintainedb y editorsf ollowingr eviewc riteriae stablished for the AGU publicationpsr ogramP. ublicationo f the Seriesis aidedb y a grantf rom the National ScienceF oundationw, hichs upportms ucho f the underlyingf ield work. Prioritiesf or publication are setb y theB oardo f AssociatEe ditorsI.n quiriesa boutp ublishevdo lumesw, ork in progresso r new proposalms ay be sentt o AntarcticR esearchS eries,A GU, 2000 FloridaA venueN W, WashingtonD, C 20009 (http:/www.agu.orgo),r to a membero f the Board. BOARD OF ASSOCIATE EDITORS RodneyM . Feldmann,C hairman,P aleontology JohnB . Anderson,M arine Geologya nd Geophysics Robert A. Bindschadler,G laciology David H. Bromwich.,M eteorologya nd UpperA tmosphereP hysics Nelia W. Dunbar, Geology StanleyS . JacobsO, ceanography JerryD . Kudenov, Marine/PolychaetBei ology JohnC . Priscu, TerrestrialB iology xi PREFACE The Long-TermE cologicaRl esearch(L TER) Pro- anda ppropriatfeo r long-terms tudies. gram was establisheidn 1981b y the UnitedS tates The primarilym arinef ocuso f thisv olumer eflects National ScienceF oundationin recognitiono f the need differencesin habitata rea and speciesri chnessb etween to studye cologicparl ocessoevse rt imes panlso ngetrh an the marinea ndt errestriael cosystemisn Antarctica.T he thoseo f mostr esearchg rantsT. he LTER Networkn ow geographicbarl eadtho f the volumeis restrictetdo the consistos f 18 sitest hats pana varietyo f ecosystemty pes westernA ntarcticP eninsulare gion,b ut the emphasiso n in the continentalU nited States, Alaska, Puerto Rico, synthesiesx tendsth e deptha ndm ultidisciplinanrya ture and Antarctica. The addition of the Palmer LTER at of thec overageT. hus,g eographicarlleys tricteodb serva- PalmerS tationA, ntarcticain October1 990e xtendedth e tions of patternsa nd distributionasr e placedi n the geographiacnald e cologicraaln geo f theL TERN etwork contexot f presenkt nowledgoef theu nderlyinpgr ocesses andp rovidetdh eo pportunittoy l inkp hysicaaln de colog- that affect them, and this knowledgec omesf rom many icalp rocessiens t hen ortherann ds outherhne mispheresr.e gionsi n the SouthernO cean. One criterion for establishmento f a LTER site is the The first paperi n thisv olumep rovidesa historical existence of historical data and observationst hat can perspectivoef expeditionisn,t ernationaanl di ndividual providae b asifso rg uidinrge searcphr ogramTsh. ust,h e researcphr ogramsa,n dn on-sponsorperdo gramsT.h ese intento f thisv olumei s to bringt ogethemr eteorological, programhs avec ontributetdo the developmeonft the hydrographicbailo, logicaaln de cologicaolb servationscu rrenut nderstandinogf t hes tructuraen df unctioningo f madein theg eneraalr eaw esto f theA ntarctiPc eninsulat hee cosysteomf theS outherOn ceanw esto f theA ntarc- andt o discustsh ep rocesseusn derlyintgh eo bservations.t ic Peninsula.T he next sectioni ncludesd escriptionso f Thisc ompilatioonf availablde ataa ndo bservatioinns environmental features of the terrestrial and marine conjunctiownit ht hep resenutn derstandionfg p rocessesh abitatsw est of the Antarctic Peninsula,i ncludingt he providetsh e foundatiofno r long-termst udieosf the glacialm arines edimentisn fjords and their use as ecosysteimn t hew esterAn ntarctiPc eninsurlae gion. paleoenvironmenintadli catorsT.h ep apero nt het erres- The centrahl ypothesoisf the PalmerL TER is that trial andf reshwateer cosysteminsc ludesa reviewo f their interannuavla riabilityin the annuael xtento f packi ce biota and functionalp rocessesin additiont o habitat hasm ajori mpactosn a ll levelso f them arinee cosystem.d escriptionIn. thiss ectiont,h ee mphasiiss on thee ffect One of the objectiveosf the PalmerL TER is to assess of environmentalf eatures on the plant and animal thei mpacot f thisv ariabilitoyn r epresentatsivpee cieosf communities.T he abundancesa nd patternsa ssociated them arinep elagice cosysteimn t her egionw esto f the with the verticala nd horizontald istributionso f biological AntarcticP eninsulaT. he LTER studyr egioni s bounded populationwse sto f theA ntarctiPc eninsualar et hef ocus by the southerenn do f BransfielSdt raita nde xtends of the next groupo f papers.T hesec overd istributions alongshotroe s outho f MargueritBea ya ndo ffshorfero m and the processecso ntrollingp opulationosf fish, pen- the AntarcticP eninsulafo r 200 km (seem ap in Figure 1 guinsa, ndm arinem ammalsO. therp apersin thiss ection of Hofmanne t al., thisv olume)T. he northerne ndo f the providet he basisf or the process-orientfeodc usi n the studyr egionis at thes outherend geo f a gapi n Ad61ie followings ectioni,n whicht he currents tateo f knowl- penguin( Pygosceliasd eliae)p opulationws,h ich is edgeo f the factorsc ontrollingb iologicadl istributions believedto resultf rom thed ependencoef this specieso n and abundancesis evaluated. The papers on human sea ice. The offshorel imit of the region is near the disturbancceo nsidehr umani mpactsin the regionw esto f seawardl imit of Antarctick rill (Euphausias uperba) the Antarctic Peninsula,b oth terrestrial and marine. An distributionsa, nd the nearshorep ortiono f the region understandinogf human-induceddis turbancess,u cha s includesp robables pawningg roundsfo r Antarctic activitiesa roundp enguinro okeriesc, ommerciafli shing, silverfish(P leuragrammaan tarcticurnT).h e interactionsa nd oil spills,i s an importanat specot f any long-term of regionacll imatologayn dh ydrograpahnyd s eaic e,a nd ecological research program. A concludingp aper ther esultinigm pacot na ll trophicle velso f thee cosystemp rovideas summariyn tendetdo placet heP almerL TER west of the AntarcticP eninsulam, aket his region unique within the context of the various ecosystemsa nd pro- xiii