ANTARCTIC RESEARCH SERIES American Geophysical Biology of the Antarctic ANTARCTIC Volume 1 RESEARCH SERIES Biologyo f the Antarctic Seas Milton O. Lee, Editor Publishedw ith thea id of a grantf rom theN ational ScienceF oundation PUBLISHER AMERICAN GEOPHYSICAL UNION OF THE National Academy of Sciences- National ResearchC ouncil Publication No. 1190 ANTARCTIC Volume RESEARCH SERIES BIOLOGY OF THE ANTARCTIC SEAS Milton O. Lee, Editor Copyright¸ 1964b y the AmericanG eophysicaUl nion 151.5 Massachusetts Avenue, N.W. Washington,D .C. 20005 Library of CongressC atalogueC ard No. 64--60030 List Price, $10.00 Printed by Garamond/Pridemark Press,I nc. Baltimore, CONTENTS Introduction to the Antarctic Research Series Harry W. Wells ix Preface Milton O. Lee xi PrimaryO rganicP roductioinn the DrakeP assagaen d BransfielSdt rait SayedZ . El-SayedE, nriqueF . Mande!lia, nd Yukio Sugimura PrimaryP roductivituyn derS eaI ce in AntarcticW aters,1 , Concentrations and PhotosynthetAicc tivitieso f Microalgaein the Waterso f McMurdo Sound, Antarctica J. S. Bunt 13 PrimaryP roductivityu nderS ea Ice in AntarcticW aters,2 , Influence of Light and Other Factorso n PhotosynthetiAcc tivities of AntarcticM arine Microalgae J. S. Bunt 27 RespiratorMy etabolisma nd EcologicaCl haracteristicosf Some Fishes in McMurdo Sound, Antarctica Donald E. Wohlschlag 33 TemperaturRe esponseasn d TissueR espirationin AntarcticC rustacea with Particular Referenceto the Krill Euphausias uperba M. A. McWhinnie 63 Antarctic Foraminiferal Zonation Orville L. Bandy and RonaldJ . EchoIs 73 ImprovedT echniquefso r BenthicT rawlinga t DepthsG reatert han 2000 Meters Robert J. Menzies 93 Cataloguea nd Bibliographyo f Antarctic and Sub-AntarcticB enthic Marine Algae GeorgeF . Papenfuss Information for Authors INTRODUCTION TO THE ANTARCTIC RESEARCH SERIES Duringt he InternationaGl eophysicaYle ar discussionwse reh eld amongg eophysicists, biologistsg,e ologistsa,n d othersc, oncerningth e developmenotf a mediumf or the pub- licationo f extensivep apersr esultingf rom the researchw ork being donei n Antarctica. The purposeo f the AntarcticR esearchS eriesi s to providet his medium,p resenting authoritativew ork with uniformlyh igh scientifica nd editorials tandardfsr om leading scientistse ngagedi n Antarctic research. In a senset,h e seriesc ontinueas traditiond atingf rom the earliestd ayso f geographic explorationan d scientifice xpeditions(cid:127)the traditiono f thee xpeditionarvyo lumesw hich setf orthi n rich detaile verythingth at wass eena nd studied.B ut in mucho f the present Antarcticw ork onee xpeditionb lendsin to the next, and it is no longers cientifically meaningfutol separatteh em. HoweverA, ntarcticr esearcihn all disciplinehsa sa large degreeo f coherencaen d deservetsh e modernc ounterpartto the expeditionarvyo lumes of past decadesa nd centuries. Papersa ppearingin the seriesre presenotr iginalc ontributiontoso l engthyo r other- wiseu nsuitablfeo r publicationin standards cientificjo urnals.T he materalp ublished will be directedn oto nlyt o scientistasc tivelye ngageidn thew orkb utt o graduates tu- dentsa nd scientistisn closelyr elatedf ieldsa s well. The booksi n the seriesw ill serve botha sr eferencefos r thes pecialisatn da ss ourceosf informatio'nfo ra nyonev ersedin the biologicaaln dp hysicasl ciencesM. anyo f the earlyv olumews ill be cohesivceo llec- tions of researchp apersg roupeda rounda central theme. An editor for each book is drawnf rom the disciplineit representsa,s a re the reviewerso n whome ache ditorr elies. Early in 1963t he NationalS cienceF oundatiomn adea grantt o the AmericanG eo- physicaUl niont o initiatet hes eriesa, nda Boardo f AssociaEted itorsw asa ppointetdo crystallizea nd implemenitt . To representth e broadn atureo f the seriest,h e members of the Boardw erec hosenfr om all fieldso f Antarcticr esearchT. hey includeJ arvis Hadleyr,e presentignego logayn ds olidE arthg eophysicHsa;r ryW . Wellsa, eronomy andg eomagnetisAm. P; . Crary,s eisinoloagnyd g laciologGy;e orgAe . Llanob, otany andz oologyW; aldoL . Schmittm, arineb iologya nd oceanographMyo; rtonJ . Rubin, meteorologayn; dW aldoE . Smith,E xecutivSe ecretaryA, GU, ex ol(cid:127)cio. Speciaal cknowledgmeisn td uet o MiltonO . Lee,w hoh ass erveda s the editoro f the firstv olumoe f thes eriesa ndh ash elpedto seta patterno f qualityw hicht he series hopest o continuea nd enhance. AGU staff membersw ho have made contributionst o the volumein cludeM arieL . Webherw, hoh ass erveda s copye ditor,a nd CarolE . Cameronw, hoh ash andledth e production. HaaaY W. WELLS ChairmanB, oardo /,4ssociatEe ditors ,tintarctic Research Series Antarctic Research Series Biology of the Antarctic Seas Vol. 1 PREFACE This, the first volume of the Antarctic Research Series, deals with a number of marine studies. There is an abundance of living forms, plant and animal, in the cold waters o~erlyingth e continental shelf of Antarctica and in the adjacent ocean depths. There are good reasons for the ongoing and currently intensified interest in marine biological research in the region, even though the difficulties in pursuing it are great. The eco- logical systems of this large polar continent and its adjacent seas are unique and there- fore demand full study; phenomena of adaptation, acclimation, and evolution are crucial. All these factors combined to make it desirable to concentrate the publication of definitive Antarctic biological research studies in the Antarctic Research Series, rather than scattering them in divers media. This is possible at this stage, since the number of studies in progress is limited, papers of unusual hut necessary length can he accom- modated, and because the authors, despite the variety of their fields of specialization, have a community of interest in Antarctic hiolosy. M.O.L. Copyright American Geophysical Union Antarctic Research Series Biology of the Antarctic Seas Vol. 1 PRIMARY ORGANIC PRODUCTION IN THE DRAKE PASSAGE AND BRANSFIELD STRAIT SAYED Z. EL-SAYED Departmenot /Oceanographayn dM eteorology Texas.4 . andM . UniversityC, ollegeS tation ENRIQUE F. MANDELLI zlrgentineN avy Hydrographi0c1 (cid:127)ce,B uenosA ires YUKIO SU½IMURA MeteorologicaRl esearchIn stitute,T okyo,J apan Abstract. Measurementso f primary production (phytoplankton) and chlorophylla content were made in the Drake Passagea nd Bransfield Strait during February and March 1963. In terms of chlorophylal and primary productiont,h e BransfieldS trait is considerablmy ore productivet han the Drake Passage;a verageds urlYacvea lueso f chlorophylla and C(cid:127) uptake are 2.4 mg/m' (cid:127) and 8.2 mg C/h/m s, respectivelyf,o r the BransfieldS trait, comparedw ith 0.73 mg/ma and 3.5 mg C/h/m '(cid:127) for the Drake PassageT. he concentrationosf surfacec hlorophylal and C(cid:127) uptake in the Drake Passage decreasedb etweenT ierra deI Fuego and the Antarctic convergence;s outh of the convergencet hey graduallyi ncreasedr,e achingt heir highestv aluesn orth of the South ShetlandI slands (chlorophyll a, 3.5 rag/mS; C(cid:127) uptake, 14.1 mg C/h/mS). The vertical distribution of chlorophyll a from the surfacet o a depth of 50 meters showedl ittle variation in the Drake Passage,b ut concentrationsw ere highesta t 10 metersi n the BransfieldS trait. The hydrographicc onditionsa nd the distribution of the nutrient elementsi n the Drake Passagea nd the Bransfield Strait were also studied and are described in this paper. INTRODUCTION along the Argentine continentals helf, in Tierra dd Fuegow atersa, nd in the DrakeP assagaen d Bransfield AlthoughA ntarcticw atersa re recognizeads one of the Strait. The data to be discussed here are from this world's most productiveo ceanic regions,i t is only investigationw, hich is still under way; they are pri- recentlyt hat interesth as beens hownb y severali nves- marily thoset aken aboardt he Argentinev esseAl RA tigatorsin studyingp rimaryp roductionb y phytoplank- CapitanC anepad uringt he February1 5 to March 20, ton in these waters. Steemann Nielsen and Jensen 1963, cruiset o the Drake Passagea nd the Bransfield [1957] indicatedt hat the rate of productioni n Ant- Strait. Sea water samplesw ere collecteda t a number arcticw atersh as beenp erhapsa little underestimated. of stationsa long the Argentine continentals helf and Klyashtorin[ 1961] measuredp rimary productioni n in Tierra del Fuegow aters,b ut the resultso f the data Antarctic watersb y the radio-carbonm ethod during collecteadt theses tationasr en oti ncludedin thisp aper. the voyageo f the Ob' aroundt he Antarctic continent during the 1959-1960 austral summer. Burkholder METHODS and Sieburth[ !961], usingc hlorophylal as an index of photosyntheticca pacityo f the plankton,s tudiedt he At eachs tation,s urfacew ater samplesw ere collected distributiono f chlorophylal in the region betweent he witha plasticb uckefto r photosyntheasnisd c hlorophyll South Shetland Islands and the Antarctic Peninsula. a determinationSsu. bsurfacsea mplefso r pigmenat nal- Early in 1962 an investigationw as initiated to meas- ysisw erec ollectewd ith a nonmetallisca mpleor f the ure the seasonaal nd yearly variationsi n productivity Van Dorn type. Chlorophyall was measurebdy a Copyright American Geophysical Union Antarctic Research Series Biology of the Antarctic Seas Vol. 1 2 EL-SAYED3(cid:127)J ANDELLAI(cid:127)N D SUGIl(cid:127)IURA BeckmanD U spectrophotomeatcecro rdingto the Mosto f the chemicaal nd pigmenta nalysews ere methodo f Richardsa nd Thompson[1 952], as modi- made aboard the researchv essel. The radioactive fledb y Creitza ndR ichards[1 955]:P rimaryo rganic Milliporef ilter disksw erea ir-shippetdo the Texas productionw asd eterminefdo r surfaces ampleso nly, A. and M. Universityw, heret heir radioactivityw as using the CT Mm ethodo f SteemannN ielsen[ 1952]. measurebdy meanso f a nuclearC hicagom odelD 47 Waters ampleasf, teri noculatiowni th NaHCX4Owa,e re gasf lowp roportioncaol unters, calemr odel1 86. held for 4 hours in a water-cooled incubator illumi- nated with fluorescenlitg ht (1200 foot-candles)T. he incubatotre mperaturfleu ctuatesdl ightlyb ut seldom RESULTS exceedetdh ato f surfacew atera t the time of the experi- ments(2 ø-3øC)T. heC T Md atat o be discussheedr e Distrœtibotn,o ] Chlorophay lal ndC (cid:127) 4 Uptake haveb eenc orrectoendl yfo rd arkb ottluep takeS. ince DrakeP assageN.i nes tationwse reo ccupieind the a submarilnigeh tp hotomewtearsn ota vailabdleu ring DrakeP assabgeet weeTnie rrad elF uegaon dt heS outh thisc ruisen,o e fforwt asm adteo u seth ei ncubatfoorr Shetlanisdl ands(F igure1 ). Thes tationwse rea r- conductsinimg ulatiend s itue xperiments. rangetdo includthe osoec cupidedu ringth ec rossing Physicaaln dc hemicadla tao btainedin cludetde rn- and recrossinogf the PassageA. few stationtsh at peratured,i ssolveodx ygen(W inklemr ethod)s,a linity werem issedo,w ingt o a stormw, erel ater occupied (Knudsemne thodp),H ,a lkalinityin, organpich osphated uringt he recrossingT.h e distributioonf surface by a modificatioonf the methodo f Robinsowni th chlorophay lal ndC x 4u ptakein theD rakeP assagise Thompso[1n9 48],n itrate[ Muiiina ndR iley,1 955], givenin Table1 andp lotteidn Figure2 . Thisf igure nitrite[ BendschneaidnedrR obinso1n9, 52],a nds ili- clearlys howtsh att hec oncentratoiofn c hlorophyal l cares[R obinsoann dT hompso!n9,4 8]. andp rimaryp roductiodne creasebde tweeTni errad el Water samplefso r phytoplanktocno untsw eret aken Fuegoa nd latitude5 8øS, with a pronouncedd ecline by the Nansenb ottlesa t 0, 10, 20, 30, 50, 75, and 150 betweens tations3 2 and 18. South of this latitude, meters.F or collectingp hytoplanktotno be usedi n theirc oncentratiognrsa duallyin creasedre, achingh igh identifyings peciesa, Hensen-typ'Ke itahara'n et (no. valuesn ortho f theS outhS hetlanIds lands( chlorophyll 13) was used. The zooplanktonst andingc rop was a, 3.5 mg/m:(cid:127); Cx (cid:127) uptake, 14.1 mg C/h/mS). It studiedfr omc ollectionms adew ith a 1-meterp lankton is noteworthtyh att he positiono f station3 0, wheret he net (no. 3) anda 12-in.C larke-Bumpsuasm pler(n o. lowesvt alueso œc hlorophyall andp rimaryp roduction 8). The resultso f the phytoplanktoann d zooplanktonw eree ncounterecdo, incidedw ith the positiono f the studiesw ill be reportede lsewhere. Antarcticc onvergenctoe be discusseidn a later section. TABLE I. Physical, Chexnicala, nd.P roductivity Data of SurfaceW ater Sa(cid:127)nplesT aken in the l)rake .Passage and BransfieldS trait during the February-March '1963C ruise Water Photo- Tem- Ct a synthetic Position peru- Alka- Chlor- Uptake, Index, Stiotan-S L atituWdLe o ngituDdaet e (TLiomceatøu l)Cr e ,g S/k, g m0(cid:127)l,/ l0 %2/ 0(cid:127)P%gaO t.a/(cid:127)1-gP Sa t,i .,/ tlN(cid:127) gaOt .a/(cid:127)Nl-g NaO t,. a/1-pN H, mlineiqtypm/1,h g ya/ml,Cl am/h a(cid:127)g mC(cid:127)nhgCa l// mh g Drake Passage 32 55006.3' 66o05.9 ' 3-13-63 1652 8.9 32.58 6.45 101 1.23 5.7 5.4 0.27 8.10 2.32 0.$I 8.53 10.47 18 55040.5 ' 66004.9 ' 3-1-63 1846 0.5 34.05 6.78 101 1.86 9.4 14.2 0.31 8.00 2.25 0.25 0.72 2.81 19 56051.5 ' 65ø01.0 ' 3-2-63 0440 5.9 34.07 6.83 100 1.78 5.2 17.4 0.29 8.05 2.35 0.17 0.62 3.65 31 57026.6 ' 64019.0 ' 3-10-63 1310 5.0 34.14 6.90 99 1.93 5.3 8.02 2.40 0.29 1.43 4.84 80 5802'7 .6 6301'0 -8 3-10-680 316 2.2 34.01 7.85 90 2.59 15.2 2;i& 0](cid:127) 7.90 2.47 0.14 0.66 4.75 20 58052.8 ' 62'44.0' 3-2-63 2159 2.0 33.86 7.57 100 2.05 20.8 15.7 0.31 8.00 2.44 0.35 1.11 3.16 29 60ø00.0' 61ø25.0' 3-9-63 1354 1.2 33.88 7.57 100 2.00 24.7 14.9 0.22 7.97 2.32 0.44 1.61 3.65 28 60053.0 ' 60021.0 ' 3-9-63 0417 1.2 33.79 7.57 0S 1.95 28.1 18.3 0.16 7.88 2.48 0.58 3.17 5.47 27 61039.8 ' 59021.2' 3-8-63 2102 0.9 33.87 7.71 100 1.89 47.8 18.9 0.16 7.95 2.56 3.53 14.1 4.0 Bransfield Strait 2I 62035.8 ' 62ø19.2 ' 3-5-63 0345 0.9 33.74 7.49 97 1.95 27.I 19.I 0.21 8.00 2.48 0.74 2.5 3.37 22 64000.0 ' 61041.2' 3-5-63 1453 0.6 33.77 7.05 90 2.36 44.1 21.0 0.18 8.90 2.46 1.38 6.84 4.95 23 63(cid:127)43.1 ' 61020.3 (cid:127) 3-5-63 2225 --0.7 34.22 7.07 90 2.52 39.7 24.2 0.21 7.88 2.48 0.3 3.0 9.93 24 63ø27.2 ' 60031.8' 3-6-63 0828 0.5 34.21 7.52 96 2.34 69.5 24.2 0.25 7.90 2.55 1.04 4.72 4.54 25 62ø56.1' 59'19.6' 3-7-63 0918 0.6 34.23 7,68 99 2.05 67.5 18.9 0.25 7.95 2.48 5.42 14.16 2.61 26 62ø31.2' 58029.0 ' 3-7-63 1536 0.5 34.20 7.85 101 2.11 71.5 16.2 0.23 7.98 2.56 5.6 18.0 3.21 Copyright American Geophysical Union Antarctic Research Series Biology of the Antarctic Seas Vol. 1 PRIMARYO RGANICP RODUCTION 3 50 ø 28 TH...TLAND sH(cid:127)sLS- 21 sTRAIT 7'50 70 o 65 ø 60 ø 55 o Fig. 1. Position of stationso ccupiedi n the Drake Passagea nd BransfieldS trait. The vertical distributiono f chlorophylla and tem- of chlorophylla concentrationa t the various depths perature from the surfacet o a depth of 50 meters in sampleda crosst he Drake Passagef rom its northern the DrakeP assages tationsis plottedi n Figure 3. This border to the Antarcticc onvergenceM. inimum values figure showsli ttle variationi n chlorophylcl oncentra- from the surface to 50 meters were found at the con- tion with depth. Slight decreasesa re noted, however, vergence( station3 0). Southo f the convergencteh ere in chlorophylcl ontenta t 50 metersi n the southD rake was a steadyi ncreasein the chlorophylal , reachinga PassageF. igure3 alsos howsth e progressivde ecreases maximum value north of the South Shetland Islands. Copyright American Geophysical Union