TheCryosphereDiscuss.,9,6187–6222,2015 D www.the-cryosphere-discuss.net/9/6187/2015/ isc doi:10.5194/tcd-9-6187-2015 us s ©Author(s)2015.CCAttribution3.0License. io n P a Thisdiscussionpaperis/hasbeenunderreviewforthejournalTheCryosphere(TC). p e PleaserefertothecorrespondingfinalpaperinTCifavailable. r | Antarctic slush-ice algal accumulation not D is c u quantified through conventional satellite s s io n imagery: Beware the ice of March P a p e r J.L.Lieser1,M.A.J.Curran1,2,A.R.Bowie1,3,A.T.Davidson2,S.J.Doust2, | A.D.Fraser4,1,B.K.Galton-Fenzi1,2,R.A.Massom1,2,K.M.Meiners1,2, D J.Melbourne-Thomas1,2,P.A.Reid5,P.G.Strutton3,6,T.R.Vance1, is c M.Vancoppenolle7,K.J.Westwood1,2,andS.W.Wright1,2 us s io 1AntarcticClimate&EcosystemsCooperativeResearchCentre,UniversityofTasmania, n P PrivateBag80,7001Hobart,Australia a p 2AustralianAntarcticDivision,ChannelHighway,7050Kingston,Tasmania,Australia e r 3InstituteforMarineandAntarcticStudies,UniversityofTasmania,PrivateBag129, | 7001Hobart,Tasmania,Australia 4InstituteofLowTemperatureScience,HokkaidoUniversity,Sapporo, D is Hokkaido,060-0819,Japan c 5BureauofMeteorology(CAWCR),G.P.O.Box727,7001Hobart,Tasmania,Australia us s 6AustralianResearchCouncilCentreofExcellenceforClimateSystemScience, io n UniversityofTasmania,7001Hobart,Tasmania,Australia P a p e r 6187 | 7Laboratoired’OcéanographieetduClimat(ExpérimentationsetApprochesNumériques), D is L’OCÉAN–UMR7159CNRS/IRD/UPMC/MNHN,IPSLBote100,4PlaceJussieu, c u 75252ParisCEDEX05,France ss io n Received:17September2015–Accepted:27October2015–Published:11November2015 P a Correspondenceto:J.L.Lieser([email protected]) pe r PublishedbyCopernicusPublicationsonbehalfoftheEuropeanGeosciencesUnion. | D is c u s s io n P a p e r | D is c u s s io n P a p e r | D is c u s s io n P a p e r 6188 | Abstract D is c u Our current knowledge of broad-scale patterns of primary production in the Southern s s Ocean is derived from satellite ocean-colour estimates of chlorophyll a (Chl a) in the ion open ocean, typically in spring-summer. Here, we provide evidence that large-scale P a 5 intra-ice phytoplankton surface aggregation occur off the coast of Antarctica during pe r austral autumn, and that these “blooms” are largely undetected in satellite ocean- colour time series (which mask the ice-covered ocean). We present an analysis of | (i) true-colour (visible) satellite imagery in combination with (ii) conventional ocean- D is colour data, and (iii) direct sampling from a research vessel, to identify and charac- c u 10 terise a large-scale intra-ice algal occurrence off the coast of East Antarctica in early ssio autumn(March)2012.Wealsopresentevidenceoftheseautumn“blooms”inotherre- n P gions(forexample,PrincessAstridCoastin2012)andotheryears(forexample,Terra a p NovaBayin2015)implyingregularandwidespreadoccurrenceofthesephenomena. e r Theoccurrenceofsuchundetectedalgalaccumulationsimpliesthatthemagnitudeof | primaryproductionintheSouthernOceaniscurrentlyunderestimated. 15 D is c u 1 Introduction s s io n InAntarctica,phytoplanktonbloomsaretypicallyobservedduringspringandsummer P a inthevicinityoftheseaiceedgeasitrecedes(Arrigoetal.,2008a),andcanalsooc- p e r curwithintheseaiceitself(Massometal.,2006).Broad-scaleassessmentofprimary productionisderivedfromsatelliteestimatesofChlabasedonocean-colourobserva- | 20 tions in the open ocean. However, ice-covered regions are masked in the processing D is (Massometal.,2006;Bélangeretal.,2007). c u In autumn, phytoplankton production declines in the Antarctic sea-ice zone due to ss limitation of growth by seasonal changes in light and temperature. Declining solar el- ion evations reduce incoming photosynthetically active radiation (PAR); reduced stratifi- P 25 a p cation allows deeper mixing of phytoplankton to depths with little light; and declining e r 6189 | temperaturesallowseaicetoreform,furtherattenuatinglightenteringtheocean(e.g. D is MitchellandHolm-Hansen,1991;KangandFryxell,1993;SmithJr.etal.,2000;Arrigo c u etal.,2008b;ArrigoandLizotte,2010). ss io Using cloud-free data from NASA’s TERRA satellite MODIS (MODerate resolution n P 5 Imaging Spectrometer) true-colour bands 1, 2 and 4 (645, 859 and 555nm), we de- a tectedalarge-scaleregion( 30000km2)ofice-associatedphytoplanktonsurfaceac- pe ∼ r cumulationthatdevelopedbetween21Februaryand19March2012(intheearlyau- | tumn)offCapeDarnley,EastAntarctica,centredon67.5◦S,70◦E(Fig.1a,apolynya D region during winter). The phytoplankton accumulation was not detected in conven- is 10 tionalmapsofChlafromtheMODISocean-colourbands(443,488and551nm).This cus intense early autumn phytoplankton occurrence, which we suggest was caused by s io auniquecombinationofphysicalandbiologicalprocesses,challengestheassumption n P thatprimaryproductionisnegligibleinthisregionduringearlyautumn(SmithJr.etal., a p 2000). While we were able to perform some opportunistic sampling, this was insuffi- er cient to conclusively establish whether this was a conventional phytoplankton bloom. 15 | However, such was the extraordinary concentration of phytoplankton within the sea- D icematrix,asobservedfromspaceandonthesurface,thatweusethetermintra-ice is c “bloom”. u s s AutumnbloomsinconsolidatedseaicehavebeenobservedaroundAntarcticapre- io n 20 viously (Fritsen et al., 1994), including in the interior parts of ice floes in the pack ice P a zone(Meinersetal.,2012).However,thesebloomswererelativelysmallinscaleand p e of low intensity. Increased autumn phytoplankton biomass has been observed in as- r sociationwithcoastalpolynyasaroundEastAntarctica(Arrigoetal.,2015;Arrigoand | vanDijken,2003)andhasbeeninferredfromicecorerecords(Curranetal.,2002)or D hypothesisedfrommodelsofice-algalproduction(seeforexampleAppendixA).Late is 25 c u summer blooms have also been reported from Antarctic waters (e.g. Comiso et al., s s 1990; Smith Jr. and Nelson, 1990), but only where the recent disappearance of sea io n icehasbelatedlyexposedcoastalwaterstohighlightlevels.Inthisstudy,weprovide P a first evidence that enhanced concentration of ice-associated phytoplankton occurs in p e r 6190 | autumn, and that this is not accounted for in current satellite-based Antarctic primary D is productionestimates. c u s s io n 2 Materialandmethods P a p e 2.1 Chronologyof2012ice-slushphytoplanktonevent r | DailyMODIStrue-colourimages(pseudoanimationinAppendixB)showthatinitiation 5 D oftheobservedphytoplanktonwereassociatedwithbreak-upoflandfastseaice(fast is c ice)totheeastofCapeDarnleybetween10Februaryand10March2012.Theinitial u s stagesweredetectedclosetothisdisintegratingfastice.Asthephytoplanktonoccur- sio n rence expanded in scale, it discoloured the sea ice (Fig. 2), which likely consisted of P 10 amixtureofpulverisedexistingiceandnewly-formedice(Massometal.,2006).This ap e icewastransportedbythewinds,firsttothesouth(particularlyinwesternPrydzBay) r andthentothenorth(relevantatmosphericmodeloutputcanbefoundinAppendixC) | withthechangeinwinddirectionoccurringaround27February.Wealsonoteevidence D ofwind-drivenstripsandpatchesofseaiceatthesouthernedgeofthe“bloom”,visible is c 15 assoutheast-northwestalignedstreakswestofCapeDarnleyaround12March.This uss processispresumablydrivenbykatabaticwinds. io n P 2.2 Watersampling ap e r On5March2012,sevenopportunisticseawatersamplesweretakeninthevicinityof | the “bloom” by the research and supply vessel Aurora Australis from the uncontami- D 20 nated seawater line intake system at 4.8m depth (see Fig. 1 for the ship track and is ∼ c samplinglocation).Photos(Fig.2)suggestaveryhighbiomassofice-associatedalgae u s attheseasurface.Samplingsiteswerelocatedbothwithinandjustoutsidethebloom sio n region, as identified from MODIS true-colour imagery. Samples were preserved with P 1%vol:volLugol’siodineandstoredinglassbottlesinthedarkat4◦C.Protistswere ap e identified using an inverted microscope at 400 and 640 magnification, and counted r 25 × 6191 | in 20 randomly chosen fields of view using a Whipple grid. Cell abundances were D is converted to biovolumes using the formulae of Hillebrand et al. (1999) and Cornet- c u Barthaux et al. (2007). Cell carbon values were then calculated from biovolumes us- ss io ingtheconversionsofMenden-DeuerandLessard(2000)andCornet-Barthauxetal. n P 5 (2007). a p e r 2.3 Evidenceofice-associatedphytoplanktonoccurrenceinautumnforother years | D The early autumn, ice-associated phytoplankton phenomenon documented here is isc u not an isolated event in this region. Using MODIS true-colour data, we identified s s 10 six other years since February 2000 when ice-associated early autumn blooms ion occurred in the same region: 2000, 2002, 2005, 2006, 2007, 2008 and 2012 P a p (http://lance-modis.eosdis.nasa.gov/cgi-bin/imagery/realtime.cgi). These blooms are e r generally most pronounced around 1 March 5days, suggesting a regular mecha- ± | nism,whichweexplorebelow. D is c u 15 3 Resultsanddiscussion ss io n 3.1 Biologicalcharacteristicsoftheseawatersamples P a p e Seawater samples taken on 5 March 2012 indicated differences in the abundance, r composition and size of the phytoplankton at sites inside and outside the bloom (Ta- | ble D1). Concentrations of Chla inside the bloom (2.11mgm−3 average, 3.3mgm−3 D 20 maximum)werethreetofivetimeshigherthanthoseoutsidethebloom(0.66mgm−3). isc Total protist abundance reached 2.6 106cellsdm−3 in the midst of the bloom com- us paredto1.2to1.7 106cellsdm−3 o×utside.Ice-associatedphytoplanktonabundance sio × n atthesurfacecouldhavebeenlargerthanat4.8mdepth,butunfortunatelycouldnot P a bemeasuredatthetime. p e r 6192 | Overall, our results show that the phytoplankton biomass was higher in the bloom D is due to higher densities and the large cell size of the dominant taxa. The phytoplank- c u toncommunitywasdominatedbymembersofthediatomgenusFragilariopsis,which ss io comprised56to75%ofallautotrophssampled.TheabsoluteabundanceofFragilar- n P 5 iopsis spp. differed little between sampling sites, but cells sampled within the bloom a p were larger (most 20 to 50µm long, with some up to 80µm long) compared to cells e r outside(commonly<10µmlong).ThehaptophytePhaeocystisantarcticawasthesec- | ond most abundant taxon, averaging 18% of all autotrophs by number and reaching 10 3n.o1t×va1r0y5scyeslltsedmma−ti3caaltlybowthithiciec-ecocvoevreerdbauntdthiceeir-frsemeallolcsaiztieon(s∼.2Cetollc5oµnmcenintradtiiaomnsetdeird) Discus meant they contributed little to overall protistan biomass. Phytoplankton taxa that in- s io habitseaice,namelyEntomoneissp.andPolarellaglacialis,werealsopresentatlow n P abundanceswithinthebloombutwereabsentoutside(TableD1). a p e r 3.2 Underestimatedimportanceofice-associatedphytoplanktoninautumn | The ice-associated phytoplankton in the region off Cape Darnley was not detected in D 15 is conventionalsatelliteocean-colourimages.Figure1ashowsMODISlevel2Chladata c u s intheCapeDarnleyregion,EastAntarctica,acquiredon2March201210:30UTC.The s io highest values are at the fringe of the area that was masked by sea ice and/or cloud n P cover, and scattered within that region. The MODIS true-colour composite has been a p overlaid in regions where sea ice and/or cloud cover prohibited Chla retrievals, and e 20 r thediscolouredseaiceandiceslushisobviousinFig.1a.Aship-borneimage(taken | 5March2012)oftheseaice/slushmatrixisgiveninFig.2.Inthisregion,thecombi- D nation of different products from the same MODIS instrument (Fig. 1a) demonstrates is c that satellite-derived estimates of Chla are lower than actual values. Also included in u s 25 Fig.1aistheship’strackwhilesamplingthebloomthreedayslater.Thistrackshows sio Chla calculated from underway fluorometry samples with the highest values of up to n P 3mgm−3 inside the bloom area, matching the highest values of the remote sensing ap e Chla estimates along the edges of the detection. Figure 1b demonstrates the close r 6193 | agreement between remote sensing data and samples in an along-track view where D is the satellite data are not masked by sea ice and/or clouds, and the high values of c u Chlaofupto6mgm−3insidethemaskedareaofthebloompatch.Becausetheyhave ss io notbeendetectedinthepast,large-scaleautumnbloomswithintheseaicezonehave n P 5 notbeenfactoredintocalculationsofregional-scaleprimaryproductionfortheSouth- a p ern Ocean (e.g. Arrigo et al., 2008b). These results are evidence that ice-associated e r phytoplanktonproductionisunderestimatedinotherseasonstoo(e.g.Massometal., | 2006;MassomandStammerjohn,2010). D is 3.3 Proximaldriversofbloomformationinautumn c u s s 10 Wehypothesisethatthecombinationoffiveenvironmentalvariablescreatethecondi- ion tionsthatleadtothisearlyautumnbloomformation;seaice,winds,light,nutrientavail- P a abilityandlackofgrazingpressure(seeconceptualrepresentationshowninFig.3a). pe r In early March, the interaction of these variables allows for an environmental window inwhichtheseice-associatedbloomscanform(Fig.3b),asfollows. | D 15 3.3.1 Seaiceconditions iscu s s BasedontheMODIStimeseriesandinformationinFig.2,thisearly-autumn“bloom” io n appears to be associated with the decay of fast ice and new sea-ice formation. Sea- P a icegrowthintheCapeDarnleyregiontypicallybeginsinlateFebruarytoearlyMarch p e r (Massom et al. (2013) and Fig. 3b), driven by the air-sea temperature gradient and winds. Frazil ice formation can act to concentrate phytoplankton in the newly-forming | 20 sea ice matrix, with frazil crystals “scavenging” algal cells from the water column as D is they rise (Garrison et al., 1983). One-dimensional biophysical model simulations (de- c u tailed in Appendix A) suggest that frazil scavenging (Garrison et al., 1989) is not es- ss sential to Chla accumulation in the ice, but would accelerate it. MODIS ocean-colour ion imageryindicatesthattherewaschlorophylinthesameregionpriortoinitiationofthe P 25 a p ice-associated bloom (see Fig. E1), possibly as a result of decay of fast ice. Fast ice e r 6194 | decay occurs through break-up and pulverisation by wind and wave activity. That de- D is cay releases both concentrated algal cells and nutrients to the surrounding ice slush c u (Massom et al., 2006). We suggest that concentrated phytoplankton cells, within this ss io interstitial sea ice matrix between brash ice and floes, comprise the discolouration of n P 5 seaiceobservedinMODIStrue-colourimagery(Fig.1a)andfromtheship(Fig.2). a p e r 3.3.2 Windconditions | Strong winds in the Cape Darnley region in March 2012 are indicated in the high- D resolution polar realisation ACCESS-P model output (Appendix C). Such conditions isc u (Fig. 3b) favour both the break-up and pulverisation of fast ice, and the formation of s s newandfrazilice.Windactionmayalsoplayakeyroledeliveringnutrientsbyaeolian io 10 n transport (Winton et al., 2014) from the continent, as well as increasing the spatial P a extentbydispersionofthebloomthroughoutitslifetime. pe r 3.3.3 Light | D AtthehighlatitudeatwhichthebloomwasobservedinMarch,theavailabilityofincom- is c 15 ingPARissufficienttoinitiatebloomconditionsifphytoplanktonareconfinedtoashal- us s lowice-slushmatrixonthesurface.Inthisway,theyareexposedtohigherlightlevels io n thanisthecaseinice-freeareaswheredeepermixingcanresultinlightlimitation,or P a withinconsolidatedseaicewithasnowcoverwheresnowsignificantlyattenuatesPAR p e (Massometal.,2006).Atsuchalowlightangle,seaicealsoactstotrapandscatter r thelightwhichthephytoplanktoncanuse(BuckleyandTrodahl,1987). | 20 D 3.3.4 Nutrientavailability is c u s PrimaryproductionintheSouthernOceanisstronglylimitedbytheavailabilityofnutri- sio n ents(Blainetal.,2007).Followingspringblooms,theexhaustionofnutrients,particu- P larlyiron,togetherwithincreasedgrazingpressureanddecreasedstratificationleads ap e toadeclineinproductionintheopenocean(deBaaretal.,1995). r 25 6195 | IntheregionoftheCapeDarnleybloom,thepresenceoflargediatomsisconsistent D is with iron enrichment (Coale et al., 2004). Sea ice and fast ice has been shown to c u contain high concentrations of iron relative to adjacent areas of open water (van der ss io Merweetal.,2011),soitssubsequentpulverisationbywaveslikelyreleasedironinthe n P 5 bloomregion,toinitiateormaintainalgalgrowth. a p Other potential sources of iron to support this early autumn bloom include: iceberg e r melt, wind-driven snow deposition, perhaps containing iron-rich dust, from the conti- | nent and wind-driven upwelling (see Fig. F1). Of these, iceberg sources of nutrients D aredifficulttoquantifyfortheCapeDarnleybloom,butfasticebreak-upwasobserved is 10 and upwelling can be inferred from wind strength and direction. Fast ice break-up in cus western Prydz Bay was observed in MODIS true-colour imagery ten days before the s io detectionoftheice-associatedbloom(AppendixB). n P a p 3.3.5 Grazing e r | Within the bloom region, the habitat provided by sea ice provides a protective matrix against zooplankton grazing (Krembs et al., 2000). Microbial grazers, such as het- D 15 is erotrophic flagellates, dinoflagellates and ciliates, were more abundant in the bloom, c u s buttheirgrazingwasinsufficienttolimitphytoplanktongrowth.Intheabsenceofsam- s io ples to indicate metazoan grazer densities, it is not possible to quantify total grazing n P pressurefortheice-associatedbloom. a p e r 3.4 Climateindices 20 | The discovery of this bloom and the satellite-based evidence of other such blooms in D is different years and regions, and implications for primary production estimates, raises c u the question of inter-annual variability. Our initial investigation suggests that the pos- ss itive phase of El Niño-Southern Oscillation (ENSO) and the Southern Annular Mode ion 25 (SAM)mayplayarole(seediscussioninAppendixG).Understandingthesedriversof Pa p e r 6196 | inter-annual variability, and their association with blooms of this type, is important for D is estimatingthescaleofthisadditionalsourceofprimaryproduction. c u s s io n 4 Conclusions P a p e Thisintra-icephytoplankton“bloom”appearstobetheconsequenceofacombination r 5 ofseaiceconditionsresultingfrombreakupoffasticeandinitialgrowthstagesofnew | sea ice. We suggest that the combination of sufficient light, nutrients released from D disintegrating ice, subsequent wind redistribution and a reduction in grazing pressure is c u allows for this phytoplankton bloom. Blooms of this type are a potentially important s s sourceofprimaryproductionintheSouthernOcean,whichisnotcurrentlyquantified io n 10 withocean-colourremotesensing,andthereforehaveimplicationsforSouthernOcean Pa foodwebsandbiogeochemicalcycling(Murphyetal.,2012). p e r | AppendixA: One-dimensionalbio-physicalmodelresults D is c ThemodelcouplesasimpleN-Pmodulewiththehalo-thermodynamicmodel(Vancop- u s penolleetal.,2010).Thismodelcapturesicethermodynamics,halo-dynamics,aradi- sio ation scheme, and a simple nutrient-phytoplankton ice algae model, characterised by n 15 P one species of ice algae (diatoms) and two limiting nutrients (nitrates and silicates). a p e Provided that iron is plentiful, the algal biomass observed in MODIS true-colour im- r agery and in-situ (from the research vessel Aurora Australis) originates either from | (i) in-ice growth of algae, or (ii) scavenging of algal cells from the water column by D 20 risingfrazilicecrystals.Thesealternativehypothesesweretestedusingsimplecalcu- isc lationsandmodelsimulations. us s io n P a p e r 6197 | A1 Simplecalculations D is c u A1.1 In-icegrowthoficealgae s s io n We first compute the amount of Chla that is achievable from the in-situ growth of ice P a algaebyconsumptionofthethelocallyavailablenutrients.Assumingthat30%ofthe p e 5 oceanicconcentrationofsilicatesofCSii =40mmolm−3 isstoredintheseaiceduring r formation(therestbeingrejectedbybrinegravitydrainage)andfullyexhaustedbyice | algae,thenwegetthatchlorophyllaconcentrationinseaiceis: D is c CCi hla=rCChlarSCiCSii×30%≈17mgChlam−3 (A1) ussio n where rCChla=0.15mgmmol−1 is a standard chlorophyll:carbon ratio in sea ice di- Pa atomsandrC=9.09istheC:Simolarratio(Sarthouetal.,2005). pe 10 Si r A1.2 Scavengingoffrazilcrystals | D TheCapeDarnleypost-polynyaishighlyproductiveinlatesummer(seeArrigoandvan isc Dijken,2003),withsurfaceChlaconcentrationsreachingupto5mgChlam−3.Assum- uss ingthatafractionα=10%ofseawaterChlawithaconcentrationCCohla=1mgm−3 is ion 15 scavenged by raising frazil crystals over an oceanic depth of ho=50m, the resulting Pap averageChlaconcentrationin30cmthickseaiceshouldbe: e r αCChlah | CCi hla= ohi o ≈17mgChlam−3. (A2) Dis c u Thesetwoscenariosgiveequivalent(high)Chlaconcentrationsanddonotdistinguish ss io betweenourtwohypothesesforthebloomsource. n P a p e r 6198 | A2 Modelsimulations D is c u Themodelisdesignedtorepresentun-deformedlevel(columnar)ice,weapplyithere s s totheseaicemixturethatcharacterisedtheCapeDarnleyearlyautumnbloom.Thisis io n basedontheassumptionthatnutrientpathwaysinlevelice(e.g.brinechannels)that P a 5 arecapturedintheone-dimensionalmodel,behavesimilarlytothefrazilmixturefound pe r betweenpancakes(althoughthismixturelikelyoccupiesmorevolumethanbrinechan- nels,becauseoftheconstantswellforcingthattendstopreventfulliceconsolidation). | AsChlaisaparticulatetracer,weassume100%ofincorporation(CChla=CChla).The D i o is modelthereforearguablyprovidesarough,minimal,representationoftheCapeDarn- cu s leybloomconditions. s 10 io n Forcingandinitialconditions Pa p e r Theseaicemodelisforcedatboundarieswiththefollowingnumbers: | – seasurfacesalinity=34gkg−1; D – nitrate concentration in seawater = 30mmolm−3 (typical value from that region, iscu 15 seeSarmientoandGruber,2006); ssio – silicateconcentrationinseawater=40mmolm−3 (typicalvaluefromthatregion, nP a seeSarmientoandGruber,2006); p e r – atmosphericheatfluxesfromempiricalformulaforsensible,latent,shortwaveand | long wave fluxes, fed by daily NCEP temperatures, winds and specific humidity D 20 (multipliedby0.83tocorrectaknownsystematicbias),togetherwithaclimatology is c ofcloudfraction; u s s – aprescribedverysmallsnowfallrate. ion P Modelrunsstartatday-of-year50(19February)whenthesurfaceenergybudgetbe- ap e comes negatives, which (approximately) corresponds with the onset of ice formation. r 6199 | Runs last 30 days, until day-of-year 80 (20 March), and are initialised with standard D is thinicecharacteristics: c u s s – icethickness=5cm; io n P – snowdepth=2cm; a p e 5 – icesalinity=22.93gkg−1(fromtheformulaofKovacs(1996)givingbulkicesalin- r ityforyoungiceasafunctionofitsthickness); | D – nitrateconcentration=20.23mmolNm−3 (sameice-oceanratioasforsalt); is c u – silicateconcentration=26.97mmolSim−3 (sameice-oceanratioasforsalt). ssio n TheconcentrationofcarbonstoredinicealgaeCDA isalsoprescribedattheoceanic P i a 10 boundary(CDoA).Asthisconcentrationintheoceanisusedasaninitialconcentration per informingnewice,accretedattheicebase,wemadeseveralsensitivityexperiments. | – Exp.1:CDoA=0.5mmolCm−3;µ=1.3×10−5s−1; D is – Exp.2:CDoA=0.5mmolCm−3;µ=1.95×10−5s−1; cus s – Exp.3:CDoA=5mmolCm−3;µ=1.3×10−5s−1; ionP 15 – Exp.4:CDoA=50mmolCm−3;µ=1.3×10−5s−1, aper whereµisthemaximumspecificphotosynthesisrate. | Experiment1correspondsto<0.1mgChla m−3(nobiologicalactivityintheocean). D In experiment 2, we tested the role of the maximum specific growth rate µ, which is c is rather uncertain. It would be bigger if the diatoms are adapted to high-light levels u s 20 (ocean diatoms) and lower if they are adapted to low-light levels (ice diatoms). In ex- sio n periment 3, the seawater that sea ice is forming from is typical of bloom conditions P ( 1mgChla m−3), but there is no harvesting of organic matter by frazil crystals. In ap ≈ e experiment 4, the seawater is typical of bloom conditions, and we assume that initial r 6200 | concentration in forming ice is enhanced by frazil scavenging (initial concentration of D Chlainseaice≈10mgChla m−3). iscus s A3 Results io n P a The model grows 50cm of ice thermodynamically within the 30day simulation period p e (Fig. A1). The cold air temperatures imply low temperatures and high brine salinities r 5 in the upper ice, which is improper to ice algae growth, as can be seen from the lim- | itation factors in the lower panels of the Fig. A1 (0 = improper to ice algal growth, D 1 = ice algal growth is possible). The four sensitivity experiments give quite different isc u blooms, particularly in terms of Chla concentration and timing of maximum Chla val- s s 10 ues(Fig.A2).IntegratingChlavertically(Fig.A3a)anddividingbyicethicknesstoget ion meanconcentrationsinseaice(Fig.A3b)indicatesthefollowingdifferencesbetween P a experiments: pe r – Experiment1giveslowconcentrationsthatareprobablyunrealistic; | – Experiment4(frazilscavengingexperiment)givesthefastesticealgalgrowthrate D is andthemostabundanticealgae; c 15 u s s – Experiments2and3areinbetweentheseextremes.Experiment2indicatesthat io n light-acclimateddiatomswithhigherµmightoutcompeteicealgaeinthebloom. P a Theconcentrationsinseaicearehighandthetimingofmaximumconcentrations p e (around day-of-year 60) is consistent with observations. Experiment 3 indicates r thatevenwithoutfrazilscavenging,accumulationoforganicmatterfromtheupper | 20 oceancreatesasufficientbasisforicealgalgrowth. D is c A4 Conclusion us s io n Modelsimulationssuggestthatfrazilscavengingwouldleadtorapidandintenseiceal- P galdevelopment(Experiment4).Sensitivityexperimentsalsosuggestthatsubstantial ap e activity is possible in the ice (i) with no frazil scavenging but accumulation of surface r 25 6201 | material in forming ice (Experiment 3); (ii) with no activity in the water, but with light- D is acclimateddiatomsoutcompetingicediatomsintheslush(Experiment2).Onthebasis c u oftheseresults,itisnotpossibletodistinguishbetweenalternativehypothesesforthe ss io originofalgalcellsintheice-associatedbloom. n P a p e AppendixB: AnimationofMODIStrue-colourimagery r 5 | A series of daily MODIS true-colour images is shown in this animation, available D fromftp://ftp.acecrc.org.au/pub/jllieser/MODIS_Animation.pps(9.3MB).Thesequence is c u startswiththeinitiationoftheobservedoccuranceon10February2012undfinishes s s on10March2012. io n P a p e 10 AppendixC: LinktoACCESS-Pwinds r | AtmosphericmodeloutputfromACCESS-Pmodel(Shresthaetal.,2013),withrelation D totheobservedbloomphenomenoncanbefoundat:ftp://ftp.bom.gov.au/anon/home/ is c cawcr/perm/preid/research/algal_bloom/access_p_algal_bloom.html. u s s io n AppendixD: Watersamplebiologicalcharacteristics Pa p e r Sampling sites were located within and outside the region identified from MODIS 15 true-colour imagery as experiencing a bloom. Samples of 1L were preserved with | 1% vol:vol Lugol iodine and stored in glass bottles in the dark at 4◦C. Protists were D is identified and counted using phase and Nomarski interference optics using an Olym- c u pusIX71andIX81invertedmicroscopeat400to640 magnification.Brightfieldop- ss × io 20 ticswasalsousedtodiscriminatetaxathatcontainedchloroplasts.Protistantaxawere n countedin20randomlychosenfieldsofview,exceptforhighlyabundanttaxathatwere Pa p countedinasubsetofthefieldofviewdefinedbyanocularquadrant(Whipplegrid). e r 6202 | Cell biovolumes and carbon conversion statistics were used to calculate the cell D is biomassofprotistantaxa/groups.Theaveragecellvolumeofeachprotistantaxonwas c u calculated using formulae presented by Hillebrand et al. (1999) and Cornet-Barthaux ss io et al. (2007). Cell carbon was then calculated using the volume-specific carbon con- n P 5 versionsofMenden-DeuerandLessard(2000)andCornet-Barthauxetal.(2007). a p Species composition and biomass differed significantly between sampling sites in- e r side and outside the bloom area (PERMANOVA, P =0.028 and P =0.026, respec- | tively)(Andersonetal.,2008).BasedonSIMPERanalysis(Primer-ELtd)largediatoms D 10 (apnrded>o5m0in%anbtalyseFdraognilabriioompsaisss)e(Cxplalarkineeadn∼d4G0o%rleoy,f2th0e06d)iff.eTrheunsc,ethbeasoebdseornveadbupnrodtaisntacen iscus communitywithinthebloomareadifferedmarkedlyfromthecommunitiesinsurround- s io ingice-freeareasintermsofcomposition,abundanceandcellsizedistribution. n P a p e r AppendixE: OceancolourimageofCapeDarnleyregionfor11February2012 | WithFig.E1weprovideaMODISocean-colourscenebeforetheiceassociatedbloom D 15 (11February2012)toshowtheexistingprimaryproduction. iscu s s io n AppendixF: TruecolourimageofCapeDarnleyregionfor23February2012 P a p e FigureF1showsaMODIStrue-colourimage,illustratingpotentialiceseedingmecha- r nisms,including(1)wind-drivenformationofstripsandpatches,(2)disintegratingfast | ice,and(3)wind-blownsnowoffthefaceoftheAmeryIceShelf. D is c u s AppendixG: InteractionsbetweenENSOandSAMintheSouthernHemisphere s 20 io n Weidentifiedanassociationbetweentheyearsinwhichice-associatedearlyautumn Pa p bloomsweredetectedinMODIStrue-colourimageryforCapeDarnley,andmodesof e r 6203 | climatevariabilitythatinfluencetheSouthernOcean,specificallytheElNiño-Southern D is Oscillation(ENSO)andtheSouthernAnnularMode(SAM).TheSouthernOscillation c u Index(SOI)isanindexdescribingtheatmosphericoscillationbetweenElNiño(warm) ss io and La Niña (cool) temperature states in the tropical eastern Pacific Ocean. Vance n P 5 et al. (2013) showed that there is a circumpolar ENSO signal in the westerly winds a p around Antarctica. This strengthening was further enhanced by interaction of ENSO e r with the SAM. By austral summer in El Niño (La Niña) years, tropospheric anoma- | lies develop into a circumpolar ridge-like (trough-like) anomaly across Antarctica that D weakens (strengthens) circumpolar westerlies (Fig. G1) and interacts with the SAM is 10 (L’HeureuxandThompson,2006). cus In the Cape Darnley region, these ice-associated early autumn blooms occurred s io in years when La Niña and positive SAM conditions generated high winds (Fig. G1), n P whichisoneoftheimportantdriversidentifiedforthisbloom(seeFig.3boftheoriginal a p paper).Theshortsatellitedataset(2000–2012)limitsourabilitytoconfirmthisENSO- er SAM-bloomrelationshipinastatisticallysignificantway,butitoccursmoreoftenthan 15 | by chance (Fogt et al., 2011) and is likely linked to greenhouse gas forcing (IPCC, D 2007). Therefore, these previously unquantified blooms may increase in frequency, is c furthercontributingtoprimaryproductionintheSouthernOcean. u s s io AcircumpolarpressuresignalofENSO n P a p The interaction between ENSO and SAM during DJF is primarily the result of coinci- e 20 r dentLaNiña-SAMpositive,ElNiño-SAMnegativeorENSO-weakSAMeventsoccur- | ringmoreoftenthanbychance(Fogtetal.,2011).Otheriterations(i.e.LaNiña-SAM D negative) resulted in anomalous transient eddies in the mid-latitudes that opposed is c ratherthanreinforcedtheSAM-ENSOinteraction.DuringLaNiña-SAMpositiveevents u s 25 (Fig.G1,upperpanel),ananomalouscircumpolartroughovertheAntarcticcontinent sio andcoastalregionsispairedwithadecidedlyun-SAM-likezonalwavenumber3(ZW3) n P patternatmid-latitudes.ZW3isassociatedwithtransitionsfromstronglymeridionalto a p e stronglyzonalflow(Raphael,2007).ZW3isthoughttobeassociatedwithENSO,and r 6204 | anAustraliansectorwave 3index(AZ3–vanOmmen andMorgan,2010)isstrongly D is correlatedwithSOIduringsummer(TableG1). c u The NCEP/NCAR reanalysis (Kistler et al., 2001) was used to investigate GPH ss io anomalies. All reanalysis datasets have high latitude data problems related to data n P 5 scarcityandtrendeffectsintheSouthernHemisphere,particularlypriorto1979(Hines a p et al., 2000; Kistler et al., 2001). Data coverage improves greatly after 1979, thus we e r usedcompositesfrom1979–2009. | TheSupplementrelatedtothisarticleisavailableonlineat D doi:10.5194/tcd-9-6187-2015-supplement. isc u s s 10 Acknowledgements. We acknowledge the use of MODIS data products or imagery from io n the Land, Atmosphere Near real-time Capability for EOS (LANCE) system operated by the P NASA/GSFC/Earth Science Data and Information System (ESDIS) with funding provided by a p NASA/HQ(https://earthdata.nasa.gov).ModeldatacanbeobtainedonrequestfromM.Van- er coppenolle([email protected]).Underwayandsampledataareavail- | 15 ablefromK.Westwood([email protected]). We thank Barbara Frankel, Julie McInnes, Fanny Chever, Maurice Levasseur and Martine D is Lizotte for their work on sample processing and analysis, Nobuo Kokubun (NIPR) for in-situ c u photography,andSoKawaguchi,RobKingandTomTrullfortheirhelpfulcontributionstoini- s s tial discussions regarding the autumn “bloom”. We are grateful to science support staff and io n 20 crewaboardRSVAuroraAustralisformakingitpossibleforsamplingtooccur.Finallywethank P a SteveNicol,IanAllison,TomTrull,IanMurrayandRowanTrebilcoforconstructivecomments p e thatgreatlyimprovedanearlyversionofthemanuscript.ThisworkwassupportedbytheAus- r tralian Government’s Cooperative Research Centre program through the Antarctic Climate & | EcosystemCRC(ACECRC).JLLwassupportedunderAustralianResearchCouncil’sSpecial 25 ResearchInitiativeforAntarcticGatewayPartnership(ProjectIDSR140300001). Dis c u s s io n P a p e r 6205 | References D is c u Anderson,M.J.,Gorley,R.N.,andClarke,K.R.:PERMANOVA+forPRIMER:guidetosoft- s s wareandstatisticalmethods,Tech.rep.,Primer-ELtd.,Plymouth,2008. 6203 io n Arrigo,K.R.andLizotte,M.P.:Primaryproducersinseaice,in:SeaIce,2ndEdn.,editedby: P 5 Thomas,D.N.andDieckmann,G.S.,Wiley,NewYork,USA,283–325,2010. 6190 ap e Arrigo,K.R.andvanDijken,G.L.:Phytoplanktondynamicswithin37Antarcticcoastalpolynya r systems,J.Geophys.Res.,108,3271,doi:10.1029/2002JC001739,2003. 6190,6198 | Arrigo,K.R.,vanDijken,G.,andLong,M.:CoastalSouthernOcean:Astronganthropogenic CO sink,Geophys.Res.Lett.,35,L21602,doi:10.1029/2008GL035624,2008a. 6189 D 2 is 10 Arrigo,K.R.,vanDijken,G.L.,andBushinsky,S.:PrimaryproductionintheSouthernOcean, cu 1997–2006, J. Geophys. Res., 113, C08004, doi:10.1029/2007JC004551, 2008b. 6190, ss 6194 io n Arrigo, K. R., van Dijken, G. L., and Strong, A. L.: Environmental controls of marine P a productivity hot spots around Antarctica, J. Geophys. Res.-Oceans, 120, 5545–5565, p e 15 doi:10.1002/2015JC010888,2015. 6190 r Bélanger, S., Ehn, J. K., and Babin, M.: Impact of sea ice on the retrieval of water-leaving | reflectance,chlorophyllaconcentrationandinherentopticalpropertiesfromsatelliteocean D colordata,RemoteSens.Environ.,111,51–68,doi:10.1016/j.rse.2007.03.013,2007. 6189 is Blain,S.,Quéguiner,B.,Armand,L.,Belviso,S.,Bombled,B.,Bopp,L.,Bowie,A.,Brunet,C., cu s 20 Brussaard, C., Carlotti, F., Christaki, U., Corbière, A., Durand, I., Ebersbach, F., Fuda, J.- sio L., Garcia, N., Gerringa, L., Griffiths, B., Guigue, C., Guillerm, C., Jacquet, S., Jean- n del, C., Laan, P., Lefèvre, D., Lo Monaco, C., Malits, A., Mosseri, J., Obernosterer, I., P a Park,Y.-H.,Picheral,M.,Pondaven,P.,Remenyi,T.,Sandroni,V.,Sarthou,G.,Savoye,N., p e Scouarnec, L., Souhaut, M., Thuiller, D., Timmermans, K., Trull, T., Uitz, J., van Beek, P., r 25 Veldhuis, M., Vincent, D., Viollier, E., Vong, L., and Wagener, T.: Effect of natural iron | fertilization on carbon sequestration in the Southern Ocean, Nature, 446, 1070–1074, D doi:10.1038/nature05700,2007. 6195 is c Buckley,R.G.andTrodahl,H.J.:Scatteringandabsorptionofvisiblelightbyseaice,Nature, u s 326,867–869,1987. 6195 s io 30 Clarke,K.R.andGorley,R.N.:PRIMERV6:UserManual/Tutorial,Primer-ELtd.,Plymouth, n P UK,2006. 6203 a p e r 6206 |
Description: