Antarctic Ecosystems: An Extreme Environment in a Changing World

575 Pages·2012·12.8 MB·English

ANTARCTIC ECOSYSTEMS
Antarctic
Ecosystems
An Extreme Environment
in a Changing World
Edited by
Alex D. Rogers, Nadine M. Johnston,
Eugene J. Murphy and Andrew Clarke
OriginatingfromaThemeIssuepublishedinPhilosophicalTransactionsoftheRoyal
SocietyB:BiologicalSciences.
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Antarcticecosystems:anextremeenvironmentinachangingworld/editedbyAlexD.Rogers...[etal.].
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Numbers1477/January29,2007and1488/December29,2007)buthasbeenmateriallychangedandupdated.”
Includesindex.
ISBN978-1-4051-9840-0(cloth)
1. Ecology–Antarctica.2. Bioticcommunities–Antarctica. I.Rogers,Alex,1968-
QH84.2.A5822012
577.099809–dc23
2011037209
AcataloguerecordforthisbookisavailablefromtheBritishLibrary.
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1 2012
CONTENTS
Contributors,xi 2GLOBALCHANGEINALOWDIVERSITY
TERRESTRIALECOSYSTEM:THE
INTRODUCTION:ANTARCTICECOLOGY MCMURDODRYVALLEYS,44
INACHANGINGWORLD,1 DianaH.Wall
AndrewClarke,NadineM.Johnston,EugeneJ.Murphy 2.1 Introduction,44
andAlexD.Rogers 2.2 TheMcMurdodryvalleyregion,46
Introduction,1 2.3 Above–belowgroundinteractions,46
Climatechange,2 2.4 Thefunctioningoflowdiversitysystems,50
Thehistoricalcontext,3 2.5 Effectsofglobalchangesoncoupled
Theimportanceofscale,3 above–belowgroundsubsystems,51
Fisheriesandconservation,4 2.6 Temperaturechange:warming,52
Concludingremarks,6 2.7 Temperaturechange:cooling,54
References,6 2.8 Directhumaninfluence:trampling,54
2.9 UVRadiation,55
PART1TERRESTRIALANDFRESHWATER 2.10 Concludingremarks,56
HABITATS,11 Acknowledgements,56
References,56
1SPATIALANDTEMPORALVARIABILITY
INTERRESTRIALANTARCTIC 3ANTARCTICLAKESASMODELSFORTHE
BIODIVERSITY,13 STUDYOFMICROBIALBIODIVERSITY,
StevenL.ChownandPeterConvey BIOGEOGRAPHYANDEVOLUTION,63
1.1 Introduction,13 DavidA.PearceandJohannaLaybourn-Parry
1.2 Variationacrossspace,16 3.1 Thevarietyofantarcticlaketypes,63
1.2.1 Individualandpopulationlevels,16 3.2 Thephysicalandchemicallakeenvironment,66
1.2.2 Specieslevel,18 3.3 Themicrobialdiversityofantarcticlakes,66
1.2.3 Assemblageandecosystemlevels,20 3.3.1 MethodsforexploringAntarcticlake
1.3 Variationthroughtime,25 biodiversity,67
1.3.1 Individuallevel,26 3.3.2 Microbialgroups,69
1.3.2 Populationlevel,27 3.3.3 Protists,70
1.3.3 Specieslevel,29 3.3.4 Crustacea,72
1.3.4 Assemblageandecosystem 3.4 Biogeography,74
levels,29 3.4.1 Spatialvariationandtheglobalubiquity
1.4 Conclusionsandimplications,30 hypothesis,74
Acknowledgments,31 3.4.2 Temporalvariationand
References,31 palaeolimnology,75
vi Contents
3.5 Evolution,76 5.2.1 Surfaceairtemperature,123
3.5.1 Prokaryotephysiology,76 5.2.2 Seaice,123
3.5.2 Eukaryotephysiology,77 5.2.3 Climateco-variability,125
3.6 Futureperspectives,78 5.3 Physicaloceanography,127
3.7 Acknowledgement,78 5.4 Nutrientsandcarbon,130
References,78 5.4.1 NutrientsandUCDWintrusions,130
5.4.2 Carboncycle,131
5.4.3 Dissolvedorganiccarbon,132
PART2MARINEHABITATSANDREGIONS,91 5.4.4 Sedimentationandexport,133
5.5 Phytoplanktondynamics,134
4THEIMPACTOFREGIONALCLIMATE 5.5.1 Seasonalscaledynamics,134
CHANGEONTHEMARINEECOSYSTEM 5.5.2 Roleoflight,134
OFTHEWESTERNANTARCTIC 5.5.3 Roleofnutrients,136
PENINSULA,93 5.5.4 Annualvariabilityin
AndrewClarke,DavidK.A.Barnes,ThomasJ.Bracegirdle, phytoplankton,137
HughW.Ducklow,JohnC.King,MichaelP.Meredith, 5.6 Microbialecology,138
EugeneJ.MurphyandLloydS.Peck 5.7 Zooplankton,140
4.1 Introduction,93 5.7.1 Communitycomposition
4.1.1 Theoceanographicsetting,96 anddistribution,140
4.1.2 Thehistoricalcontext,97 5.7.2 Long-termtrendsand
4.2 Predictedenvironmentalchangesalongthe climateconnections,142
westernantarcticpeninsula,98 5.7.3 Grazingandbiogeochemical
4.3 Environmentalvariabilityandecological cycling,142
response,100 5.8 Penguins,143
4.3.1 Bioticresponsestoclimatechange: 5.8.1 Contaminantsinpenguins,145
somegeneralpoints,102 5.9 Marinemammals,146
4.4 Responsesofindividualmarinespeciestoclimate 5.10 Synthesis:foodwebsofthewap,147
change,102 5.11 Conclusions,148
4.4.1 Acclimationandevolutionary Acknowledgements,149
responsestoenvironmentalchange References,149
inantarcticmarineorganisms,104
4.5 Communitylevelresponsestoclimate 6SPATIALANDTEMPORALOPERATION
change,106 OFTHESCOTIASEAECOSYSTEM,160
4.6 Ecosystemlevelresponsestoclimate E.J.Murphy,J.L.Watkins,P.N.Trathan,K.Reid,
change,107 M.P.Meredith,S.L.Hill,S.E.Thorpe,N.M.Johnston,
4.7 Whatbiologicalchangeshavebeenobserved A.Clarke,G.A.Tarling,M.A.Collins,J.Forcada,
todate?,109 A.Atkinson,P.Ward,I.J.Staniland,D.W.Pond,
4.8 Concludingremarks,110 R.A.Cavanagh,R.S.Shreeve,R.E.Korb,M.J.Whitehouse,
Acknowledgements,110 P.G.Rodhouse,P.Enderlein,A.G.Hirst,A.R.Martin,
References,111 D.R.Briggs,N.J.CunninghamandA.H.Fleming
6.1 Introduction,160
5THEMARINESYSTEMOFTHEWESTERN 6.2 Oceanographyandseaice,163
ANTARCTICPENINSULA,121 6.2.1 Upper-oceancirculationand
HughDucklow,AndrewClarke,RebeccaDickhut,ScottC. characteristicsintheScotiaSea,163
Doney,HeidiGeisz,KuanHuang,DouglasG.Martinson, 6.2.2 Physicalvariabilityandlong-term
MichaelP.Meredith,HollyV.Moeller,MartinMontes- change,167
Hugo,OscarSchofield,SharonE.Stammerjohn, 6.3 Nutrientandplanktondynamics,168
DebbieSteinbergandWilliamFraser 6.4 Krillinthescotiaseafoodweb,171
5.1 Introduction,121 6.4.1 Krilldistributioninthe
5.2 Climateandice,123 ScotiaSea,171
Contents vii
6.4.2 Krillgrowthandageinthe 8.2 Theregion,245
ScotiaSea,173 8.2.1 Theeast(80(cid:2) E–150(cid:2) E),245
6.4.3 Krillreproductionandrecruitment 8.2.2 Thewest(30(cid:2) E–80(cid:2) E),247
intheScotiaSea,174 8.3 Ecosystemchangeoffeastantarctica,251
6.4.4 Krill–habitatinteractionsinthe Summary,251
ScotiaSea,177 References,252
6.4.5 Krillpopulationvariabilityand
changeintheScotiaSea,180 9THEDYNAMICMOSAIC,255
6.4.6 KrillintheScotiaSeafoodweb,183 DavidK.A.BarnesandKathleenE.Conlan
6.5 Foodweboperation,184 9.1 Introduction,255
6.5.1 Trophiclinks,184 9.2 Historicalandgeographicperspectives,256
6.5.2 Spatialoperationofthefoodweb,189 9.3 Disturbance,257
6.6 Ecosystemvariabilityandlong-term 9.3.1 Iceeffects,258
change,192 9.3.2 Asteroidimpacts,260
6.7 Concludingcomments,195 9.3.3 Sedimentinstabilityand
Summary,196 hypoxia,261
Acknowledgements,197 9.3.4 Windandwaveaction,261
References,197 9.3.5 Pollution,262
9.3.6 UVirradiation,263
7THEROSSSEACONTINENTALSHELF: 9.3.7 Volcaniceruptions,263
REGIONALBIOGEOCHEMICALCYCLES, 9.3.8 Trawling,264
TROPHICINTERACTIONS,AND 9.3.9 Non-indigenousspecies(NIS),264
POTENTIALFUTURECHANGES,213 9.3.10 Freshwater,265
WalkerO.Smith,Jr.,DavidG.Ainley, 9.3.11 Temperaturestress,265
RiccardoCattaneo-ViettiandEileenE.Hofmann 9.3.12 Biologicalagentsofphysical
7.1 Introduction,213 disturbance,266
7.2 Physicalsetting,214 9.4 Colonisatonofantarcticsea-beds,266
7.3 Biologicalsetting,219 9.4.1 Larvalabundance,266
7.3.1 Lowertrophiclevels,219 9.4.2 Hardsubstrata,266
7.3.2 Mid-trophiclevels,225 9.4.3 Softsediments,269
7.3.3 Fishesandmobilepredators,226 9.5 Implicationsofclimatechange,276
7.3.4 Uppertrophiclevels,227 9.6 Conclusion,279
7.3.5 Benthos,229 Acknowledgements,280
7.4 Foodwebandbioticinteractions,230 References,281
7.5 Conclusions,232
7.5.1 UniquenessoftheRossSea,232 10SOUTHERNOCEANDEEPBENTHIC
7.5.2 Potentialimpactsofclimate BIODIVERSITY,291
change,233 A.Brandt,C.DeBroyer,B.Ebbe,K.E.Ellingsen,
7.5.3 Conservationandtheroleof A.J.Gooday,D.Janussen,S.Kaiser,K.Linse,M.Schueller,
commercialfishingactivityinthe M.R.A.Thomson,P.A.TylerandA.Vanreusel
RossSea,234 10.1 Introduction,291
7.5.4 Researchneedsandfuture 10.2 Historyofantarcticbiodiversity
directions,235 work,293
Acknowledgements,235 10.3 Geologicalhistoryandevolutionofthe
References,235 antarctic,294
10.3.1 IndianOcean,294
8PELAGICECOSYSTEMSINTHEWATERS 10.3.2 SouthAtlantic,294
OFFEASTANTARCTICA(30(cid:2) E–150(cid:2) E),243 10.3.3 WeddellSea,295
StephenNicolandBenRaymond 10.3.4 DrakePassageandScotia
8.1 Introduction,243 Sea,296
viii Contents
10.4 Benthiccompositionanddiversityof 11ENVIRONMENTALFORCINGAND
meio-,macro-andmegabenthos,296 SOUTHERNOCEANMARINEPREDATOR
10.4.1 Meiofauna,297 POPULATIONS,335
10.4.2 Macrofaunalcomposition PhilN.Trathan,JaumeForcadaandEugeneJ.Murphy
anddiversity,299 11.1 Climatechange:recent,rapid,regional
10.4.3 Megafaunalcomposition warming,335
anddiversity,304 11.2 Usingoscillatoryclimatesignalstopredict
10.5 Phylogeneticrelationshipsofselected futurechangeinbiologicalcommunities,337
taxa,308 11.3 Potentialforregionalimpactsonthe
10.5.1 Foraminifera,308 biosphere,338
10.5.2 Isopoda,308 11.4 Confoundingisuesinidentifyingabiological
10.5.3 Tanaidacea,309 signal,339
10.5.4 Bivalvia,310 11.5 Regionalecosystemresponsesasa
10.5.5 Polychaeta,310 consequenceofvariationinregionalfood
10.5.6 Cephalopoda,310 webs,340
10.6 Biogeographyandendemism,311 11.6 Wherebiologicalsignalswillbemost
10.6.1 Porifera,311 apparent,340
10.6.2 Foraminifera,311 11.7 Thesouthwestatlantic,341
10.6.3 Metazoanmeiofauna,311 11.8 Theindianocean,344
10.6.4 Peracarida,312 11.9 Thepacificocean,345
10.6.5 Mollusca,312 11.10 Similaritiesbetweentheatlantic,indian
10.6.6 Echinodermata,313 andpacificoceans,346
10.6.7 Brachiopoda,313 11.11 WhatENSOcantellus,347
10.6.8 Polychaeta,313 11.12 Futurescenarios,349
10.6.9 Bryozoa,313 References,349
10.7 Relationshipofselectedfaunal
assemblagestoenvironmental
variables,313 PART3MOLECULARADAPTATIONS
10.7.1 Large-scalepatternswith ANDEVOLUTION,355
depth,313
10.7.2 Patternsinfluencedbyother 12MOLECULARECOPHYSIOLOGYOF
environmentalorphysical ANTARCTICNOTOTHENIOIDFISHES(cid:3),357
factors,317 C.-H.ChristinaChengandH.WilliamDetrichIII
10.7.3 Isopoda,318 12.1 Introduction,357
10.8 Similaritiesanddifferencesbetween 12.2 Survivingthebigchill–notothenioid
antarcticandotherdeep-sea freezingavoidancebyantifreeze
systems,318 proteins,358
10.8.1 Theenvironment,318 12.2.1 FreezingchallengeinfrigidAntarctic
10.8.2 Adirectcomparisonbetweenthe marineenvironment,358
deepseaoftheSOandtheWorld 12.2.2 Historicalparadigmofteleostfreezing
Ocean,319 avoidance,360
10.8.3 Dispersalandrecruitmentbetween 12.2.3 ParadigmshiftI:the‘larval
theSOandtherestofthe paradox’,360
world,320 12.2.4 ParadigmshiftII:liverisnotthe
10.8.4 Thespecialcaseof sourceofbloodAFGPin
chemosynthetically-driven notothenioids,362
deep-seasystems,320 12.2.5 Gutversusblood–importanceof
10.9 Conclusions,321 intestinalfreezeavoidance,363
Acknowledgements,321 12.2.6 Non-hepaticsourceofplasma
References,323 AFGP,364
Contents ix
12.2.7 Alterationsinenvironmentsand 13.2.5 Ecologicalimplications,399
dynamicevolutionarychange 13.2.6 Integrationofphenomena:concepts,
innotothenioidAFGPgene resultsandperspectives,405
families,364 Acknowledgements,409
12.2.8 Summarycomments–antifreeze References,409
proteingaininAntarctic
notothenioidfish,367 14EVOLUTIONANDBIODIVERSITY
12.3 Haemoproteinlossandcardiovascular OFANTARCTICORGANISMS,417
adaptationinicefishes–dr.notothe AlexD.Rogers
rescue?,367 14.1 Introduction,417
12.3.1 Vertebrateswithouthaemoglobins– 14.2 Theantarcticbiota,418
youmustbekidding!,367 14.3 Thebreak-upofgondwanaandthe
12.3.2 Haemoproteinlossinicefishes:an evolutionofthesouthernhemisphere
evolutionaryperspective,368 biota,420
12.3.3 Cellularcorrelatesof 14.3.1 Vicarianceversusdispersal,420
haemoproteinloss,370 14.3.2 Dispersalmechanisms,421
12.3.4 Theicefishcardiovascular 14.4 Theevolutionandbiodiversityoftheterrestrial
system,371 sub-antarcticandantarcticbiota,423
12.3.5 Compensatoryadjustment 14.4.1 Plants,423
oftheicefishcardiovascularsystem 14.4.2 Animals,427
inaregimeofreducedinterspecific 14.5 Themarineenvironment,432
competition?EnterDr.NO,371 14.5.1 Biogeographyandmacroevolution,
12.3.6 Haemoproteins,NOmetabolism, 432
andicefishevolution,372 14.5.2 Notothenioidfish,432
12.3.7 Icefishesanderythropoieticgene 14.5.3 Birds,435
discovery,372 14.5.4 Marineinvertebrates,436
12.3.8 Summarycomments:haemoprotein 14.5.5 Themolecularecologyand
lossinAntarcticicefishes,374 phylogeographyofthemarinebiota,
12.4 Concludingremarks,374 437
Acknowledgements,374 14.5.6 Patternsofgeneticvariation
Dedication,374 inmarinespecies,448
References,374 14.6 Antarctica:aclimaticcrucible
ofevolution,450
13MECHANISMSDEFININGTHERMALLIMITS 14.7 Thehistoricalconstraintsonadaptation
ANDADAPTATIONINMARINEECTOTHERMS: topresentclimatechange,453
ANINTEGRATIVEVIEW,379 14.8 Futuredirectionsforresearch,453
HansO.Po€rtner,LloydS.PeckandGeorgeN.Somero References,454
13.1 Introduction:climate-dependentevolution
ofantarcticfauna,379
13.2 Phenomenaofthermalspecialization PART4CONSERVATIONANDMANAGEMENT
andlimitation,382 ASPECTS,469
13.2.1 Molecularandmembrane
aspects,383 15BIOGEOGRAPHYANDREGIONAL
13.2.2 Genomicaspects:geneexpression CLASSIFICATIONSOFANTARCTICA,471
andlossofgeneticinformation,390 P.Convey,D.K.A.Barnes,H.J.Griffiths,S.M.Grant,
13.2.3 Frommoleculartosystemicaspects: K.LinseandD.N.Thomas
thermallimitation,393 15.1 Introduction,471
13.2.4 Frommoleculartosystemicaspects: 15.2 Historicalbackground,474
thermaladaptationofperformance 15.2.1 Physicalregionsinthemarine
capacity,397 environment,474
x Contents
15.2.2 Smaller-scaleregionalization 16.2 Legalframeworksforconservation
withintheAntarcticmarine andmanagement,495
environment,474 16.2.1 Earlyregulationofmarineliving
15.2.3 Physicalregionsinthelittoral resourceharvesting,495
environment,475 16.2.2 TheAntarcticTreatySystem,497
15.2.4 Physicalregionsintheterrestrial 16.2.3 Other(non-ATS)agreements
environment,475 andtoolsrelevanttoconservation
15.3 Dataavailability,476 andmanagement,500
15.4 Differentrealmsinthemarineandterrestrial 16.3 Conservationandmanagement
environments,477 measures,502
15.4.1 Pelagicrealm,477 16.3.1 Pollutionandlocal
15.4.2 Seaice,478 disturbance,502
15.4.3 Benthicrealm,479 16.3.2 Biosecurityandnon-native
15.4.4 Theterrestrialenvironment,479 species,505
15.4.5 Biogeographicalpatternsinthe 16.3.3 Conservationandmanagement
terrestrialenvironment,480 ofmarinelivingresources,505
15.4.6 Biogeographicpatternsinthe 16.3.4 Conservationofotherindividual
marineenvironment,481 species,507
15.5 Overview,485 16.3.5 Protectedareas,509
Acknowledgements,486 16.4 Conservationscienceand
References,486 monitoring,512
16.5 Futurechallenges,515
16CONSERVATIONANDMANAGEMENT 16.6Conclusions,520
OFANTARCTICECOSYSTEMS,492 Acknowledgements,521
SusieM.Grant,PeteConvey,KevinA.Hughes, References,521
RichardA.PhillipsandPhilN.Trathan
16.1 Introduction,492 Index,526
Contributors
DavidG.Ainley C.-H.ChristinaCheng
H.T.HarveyandAssociates,3150AlmadenExpressway, Department of Animal Biology, University of Illinois,
SanJose,CA95118 515MorrillHall,Urbana,IL61801,USA
AngusAtkinson AndrewClarke
BritishAntarcticSurvey,NERC,HighCross,Madingley BritishAntarcticSurvey,NERC,HighCross,Madingley
Road,CambridgeCB30ET,UK Road,CambridgeCB30ET,UK
DavidK.A.Barnes MartinA.Collins
BritishAntarcticSurvey,NERC,HighCross,Madingley Government House, Stanley, Falkland Islands, South
Road,CambridgeCB30ET,UK Atlantic
ThomasJ.Bracegirdle KathleenE.Conlan
BritishAntarcticSurvey,NERC,HighCross,Madingley CanadianMuseumofNature,POBox3443,StationD,
Road,CambridgeCB30ET,UK Ottawa,OntarioK1P6P4,Canada
AngelikaBrandt PeterConvey
Biocentre Grindel and Zoological Museum, Martin- BritishAntarcticSurvey,HighCross,MadingleyRoad,
Luther-King-Platz3,20146Hamburg,Germany Cambridge,CB30ET,UK
D.R.Briggs NathanCunningham
BritishAntarcticSurvey,NERC,HighCross,Madingley BritishAntarcticSurvey,NERC,HighCross,Madingley
Road,CambridgeCB30ET,UK Road,CambridgeCB30ET,UK
RiccardoCattaneo-Vietti ClaudeDeBroyer
Dipartimento per lo Studio del Territorio e delle sue Institut Royal des Sciences Naturelles de Belgique,
Risorse,Universit(cid:2)adiGenova,16132Genova RueVautier29,1000Bruxelles,Belgium
RachelA.Cavanagh H.WilliamDetrichIII
BritishAntarcticSurvey,NERC,HighCross,Madingley Department of Biology, Northeastern University, 134
Road,CambridgeCB30ET,UK MugarHall,Boston,MA02115,USA
StevenL.Chown RebeccaDickhut
Centre for Invasion Biology, Department of Botany & VirginiaInstuteofMarineScience,CollegeofWilliam
Zoology, Stellenbosch University, Private Bag X1, andMary,GloucesterPoint,VAUSA
Matieland7602,SouthAfrica