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Polar Biosci.,14,]O 23. 2001

Variationin the diatom community under fastice near Syowa

Station,Antarctica,duringthe austral summer of 1997/98

Akira Ishikawai',Naoki Washiyama2, AtsushiTanimurai

and Mitsuo Fukuchi3
iRiculty
ofBioresources.Mie Uhiversity,1515 Kamihatna-cho, 71su5i4-8507
2Ryokuseisha Ybsuda Bldg. 15-l4 Tlyukiji
Corp., Tlsukiji 2-chome,Chuo-ku,7k)kpo104-O045
]IVationai
ofPolar
In,ytitute Research,Kaga J-chome, lrabashi-ku,7bkyo 173-8515

Abstract: Variationsinabundance and community structure of diatoms under the

fasticenear Syowa Stationwere inyestigated almost dailyduring the austral summer
of 1997198. Two periodsof high chlorophyll a concentration were observed through-
out the study: from the end of December te early January and from the rniddle to the
end of' JanuaTy. Size fractionationof chlorophylla revealed that phytoplankton
during the former period consisted mostly of organisms !argerthan 20stm and during
the latterperiod,1O-20"m. The largediatoms, Porosira pseudodentieulata and Ryeudo-
nitzschia turgiduioides, and small diatoms, F)ugilariopsis spp., were the dominant
organisms in the former and latterperiods,respectively, Melting of the fast ice
occurred in January, indicatjnga possibility that small sized diatomswere released

from the iceto the water column. Aceumulation of small diatomsin a sediment trap
fo11owed a decrease of their abundance in the w'ater column. These results indicate
that most of the icealgae detached from the ice sank direetly to the bottomduringthe
latterhalf of the austral summer.

key words: chlorophylL a, diatom comrnunity. water column, sinking, Syowa Station

Introduction

The importanceof ice algae, primary producers, in ice-coveredareas is well

as

known (Horner,1985a). Their abundance in the ice increases in spring, before

phytoplankton blooms in the water column (e.g. Horner, 1985a,b). There are reports
,

that the icealgae released from the ice seed the bloom in the water column, especially in
ice edge zones (Garrison and Buck, 1985; Smith and Nelson, 1985; Garrison et al.,

1987). However, Riebesellet al, (1991)contend that grazingand sinking are the main
fates of sea icebiota.
Studies on ice algae near Syowa Stationhave been undertaken since the late1950s
(Hoshiai and Watanabe, 1996). Near Syowa the icea]gal biornass reaches a maximum

in spring/early summer (Hoshiai, 1981a,b; Watanabe and Satoh, 1987) and the abun-
dance of phytoplankton inthe water column usually reaches itsmaximum at the efld of

January and in early February (e.g., Satoh et aL, 1986). The primary productiQn of
the phytoplanktonin the water column islower than that of the ice algae in the spring
but higherin the summer (Satoh and Watanabe, 1988; Satoh et aL, 1991).

'
E-mail: ishikawa@bio.mie-u.ac.jp

10

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Diatom community under the ice in summer 11

In this study we report almost dailychanges in the community structure of

phytoplanktonwith special reference to that of diatoms under the fasticenear Syowa
Stationduring the austral summer of 1997/98.

Materialsand methods

Sampling was conducted at St, A (69000.00'N, 39037.12'E),locatednear Syowa

Station, from 27 December, 1997 to 3 February,1998 (Fig. 1). The water depthat the
station was 40m. The thickness of the fasticeat the site was 260 and 215cm at the
start and at the end of the sampling The icein this season was
period,respectively.
thicker than isusually observed at the site by more than I m (e.g, Watanabe and Satoh, ,

1987). Water samples were collected with a 2l Niskinbottlethrough a holedrilled in

the ice. Collectionswere made almost dailyat 1m depth (daily water samples) and at
approximately weekly intervalsat 1.5, 10,20 and 35m depths(vertica] water samples)
below the ice. Temperature and salinity were measured with a CTD (Sea-Bird Elec-
tronics,Inc). The CTD operation was done almost dailyat depths shallower than
several meters and at approximately weekly intervalsthrough the water column.

44,
59i

's

69'oo'
S

Ol,

02i

Fig.1. Location of sampling station near Syowa Station.Antarctica,

Concentrationof chlorophyll a was determined for each water sample. A 200 ml

aliquot from both dailyand vertical water samples was filtered through a Whatman
GFIF glassfiberfilter with vacuum pressure of less than 1oo mmHg to givetotal Chl a.
Another aliquot (300-500 ml) of dailywater sample was filtered through a 20"m mesh
(>20pam fraction) without vacuum and the filtrate was sequentially filtered through a
Nuclepore filterwith pore size of IOptm (10-20ptmfraction)without vacuum. The
chlorophylla retained on the filters was extracted in IV,N-dimethylformamideat
-200C fbr 24h
(Suzuki and Ishimaru, 1990) and itsconcentration was determinedby
the fluorometric method of Parsons et aL (1984)with a Turner Design fiuorometer
Medel 10-AU-O05. Chlorephylla concentration of the < 10pam fraction was obtained
from the difference between the totalChl a and the sum of the ) 1Oum fraction.Size
fractionation of chlorophyll a for the vertical water samples was done using onty the

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12 A. Ishikawa, N, Washiyama, A. Tanimura and M. Fukuehi

20pam mesh (> or <20ptm fractions).

The daily water samples were also used for cell counts of diatoms and

dinofiagellates. Those sarnples were immediatelyfixedby adding borax buffered form-

concentration of 2%. A 50mt aliquot of the sample was settled in an
aldehyde at a final
Uterm6hl chamber forat least24 h;species identification
and cell counts were done with
a Nikon TE-300 inverted microscope at 100-400Xmagnification and, when necessary

for positiveidentification of species, cells were cieaned with concentrated HCI and
KMn04 mixture (Simonsen, 1974), driedin air, placedon aluminum stubs, coated with

gold, and examined with a scanning electron microscope (JEOL, JSM-T-200), The
species was identified by referring to Medlin and Priddle (1990)and Hasle and
Syvertsen(1997). Among the diatom population,the two dominant species were
selected to determinespecies succession. In this analysis, the two dominant species
almost always occupied more than 80% of the total diatom population.
Sinkingdiatoms were collected with a sediment trap moored 20m below the ice.
The sediment trap was 70.6 cm inheight;upper 40.6 cm cylinder, with a baMe at the top,

and lower 30cm funnelwith 500ml bottleat the bottom. The diameterwas 15.4cm,
making a collecting area of 186cm2. The collection was made duringthe periodfrorn
2 January to 2 February 1998. The traps were fi11ed with artificial sea water with

salinity of ca. 35g l and borax buffered formaldehydejust beforedeployment. The

i

trap was replaced at approximately weekly intervals. For species identification and

counting of diatoms, a 112SO-1!500 aliquot of the deposited matter was suspended in 10

ml of filteredsea water containing borax bufferedformaldehyde at 2%, From this
suspension, a 1 to 2ml aliquot was put on a slide glass and cells counted at 100-400×

magnification. Counts were carried out in triplicatefor each sample and the daily ee]1

flux (cells m'2 day'i) was calculated. Although the cell fluxis illustrated oniy for

dominant diatoms in this study, these species occupied most (83･-98%) of the total

diatorn cells.

Results

Temperature,salinity and density

A vertical profileof temperature showed that the thermocline appeared after 3

January and deepened to the end of January (Fig. 2A). Warm water exceeding OOC
seemed to be located from shallow to deeper layersas the thermocline deepened.
Maximum temperature was recorded (O.250C) at 1 m below the iceon 17 January.
The vertical salinity gradient was very small in the water column from the end of
December to the beginning of January (Fig.2B). Thereafter,the gradientgradually
appeared at depthsshallower than 15m and developedthroughout the water column by
the end of January, Minimum salinity was recorded (31.4PSU) at 1 m below the iceon

25 January. The trend of changes in salinity seemed to be opposite to that in

temperature, indicating that melting of the fastice occurred.

The water densitywas almost homogeneous in the water column deeper than 1 m
below the iceuntil the beginningofJanuary (Fig. 2C). A density gradientwas formed
in the surface layershal]ower than around 10m on 7 January and developedthrough the
water column by the end of January. No apparent pycnocline was observed.

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Diatom community under the ice in summer 13

Date
1997Dec.27 1998
Jan. Feb.3013
301 5 10 15 2025
159131721252933
･ N,
ki,,

(A)

AEv=-Qma

159131721252933 ,

Fig.2. Temporal variations of (A) temperature, (B)the

salinitvbottom density,oi, in water
column from 1 m below the fastice to near A
atandst.(C) from 27 Decemberthe
1997 to 3
February 1998.

Chlorophylla
Throughout the study, two periodsof high Chl a concentration were observed in
layersshallower than 20m: the first
from lateDecember to early Januaryand the second

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14 A. Ishikawa, N, Washiyama, A. Tanirnura and M. Fukuchi

from mid- to late January (Fig. 3A). Total Chla showed a trend of decreasing

graduallywith depth, Detailed changes in tota]Chl a 1 m below the ice are shown in
Fig.4A. Concentration of totalChl a increasedfrom the beginning of the study to 30
December (7.9xtgl' i),
decreasing to 1.3psgl ion 10 January. Total Chla increased
again from 14 January,reaching tltehighestvalue of 9.9ygl'' on 22 January. The
high abundance lasted for about 7 days,then decreased until the end of the study,
The fraction>20ptm accounted forrnore than 90% of the totalChl a through the
water column at the end of December but decreasedto < 10% at the end of January
(Fig, 3B), Detailed changes inrelative abundance of the fractions to the total Chl a at
1m beiow the iceare also shown in Fig. 4B. The fraction >20"m comprised 50-90%,
from the end of December to the beginning of January,decreasing laterin the season
(Fig. 4B). The 10-20"m fractionincreased, supplanting the >20pam fraction,from
the beginningof January,and became the major component of the total Chla by the

middle of January. Thereafter, this fraction comprised almost 60-90% of the total
Chl a. Throughout the study period, the fractionef > 10pam (sumof >20 and 10--20
ptm) accounted for62-99% of the total Chl a, except in a f'ew samples (30December,

Date
1997Dec. 1998
Jan. Feb.

1
5

10

15

20

25

30AE

3sZ'a-'

gy
iii
1,S)i],il
1oas
o'
eo}liiiil,ii>ll)ii

,1
10
7e 6e
90

15

20

25

30

35 (B)
Fig.3, Temporal variations of (A) total chlorophyll a concentration (sigii),
and (B) relative

abundance (%) of fractionlargerthan 20pam in the water column at St.A.

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Diatom community under the lce in summer 15

10

tt3u=)-`a99=-o-"
8

2
oF

o1OOAS

(B} ew >2e "m

sog=8 1 io-2o pam

n <lo "m

6osoca

40otz･-asca

20cr

o
27 1 10 20 30 3
Dec. Jan, Feb.
1997 1998
Fig. 4. Daily changes in (A)total chlorophylla concentration and (B)relative abundance of itssize
fractions at 1 m below the ice in the water column at St. A. Unfi11ed partsdenote no data.

and 21 and 24 January). No obvious trend in the change of relative abundance of the
fractionsmaller than 10"m was observed.

Diatom and dinoflagellate community structure

Daily change in cel] abundance of total diatoms at 1m below the ice (Fig. 5A) was
generally similar to that of total Chla at that depth. Name]y, high abundances of
''i
> 105cells l were observed befbreand after the low abundance (3 10`-9.6 10`cells
× ×

l i) periedfrom9to 13 January 1998. The abundance exceeded the order of 106cells

ifrom
l the middle to the end of January. On the other hand, abundance of the total

dinoflagellates(Fig. 5B) was always much lower,at a densityfrom O-103 cells l i, than
that of diatoms, Almost 100% of the species observed in the dinoflagellate population
were Protqperidinium spp,, which are heterotrophic. Centric diatoms were observed at

the relative abundance of 9-10% of the total diatom popuaation from 27 December to
3 January. The relative abundance decreased thereafter to <1% by 12 January.
Instead,pennate diatoms occupied nearly or totally the diatom populationsafter 7
January. In this study, 11 species of 8 genera of centric diatems were observed (Table
1), Among them, the large species, forosira pseudodenticulata (53.9ptm in average

diameter;n = 106),was almost always the dominant species at a relative abundance of

69-100% of the centric diatom population beforethe middle of January (Fig. 6A).

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16 A. Ishikawa, N. Washiyama. A. Tanimura and M, Fukuchi

lo7

io6

---.r9oe.-ts285o

leS

lo4lo4

lo3

lo2

o
27 1 10 20 30 3
Dec. Jan. Feb,
1997 1998
Fig. S. Daily changes in celi densitiesof (A) diatoms and (B)dinoflagellates
at 1 m belowthe ice in
the water column at St. A.

The pennate diatom population,in which 8 species and 3 unidentified groupsof 7 genera
were observed (Tablel), was characterized by the ]arge species, iFlseudo-nitzschia
turgiduloides (94.9
and 2.6ptm in average for the apical and transapicalaxes, respective-
ly;n 103),and the small
=
species, Ragilariopsis
curta andE cylindrus (23.5 and 2.5pm
in average for the apical and transapicalaxes, respectively; n=106) (Fig.
6B). The
]F)'agilariqpsis species were counted as a single taxon (D'agiiariopsisspp.) becausethe two
species were not distinguishable
by lightmicroseopy. ]Rgeudo-nitzschia turgiduloides
was the dominant species, at a relative abundance of 47-99%, from 27 December to 8
January and iF)ragilariopsis spp., at a relative abundance of 50-100%, duringthe rest of
the study period, except on 13 January. In this study, R turgiduloides often formed
long chains, whereas most cells of F},agilariqpsis
spp, were solitary or in short chains of
2-3 cells. Aggregates were often observed in the water samples.
As a result of analysis in relationships
between the cell densitiesof the dominant
diatoms and concentration of the >20 and 10-20um chlorophyll a fractions at 1 rn
below the ice, P. pseudodenticulata and PL turgiduloides showed significant positive
relationships with the >20ptm fraction; the correlation coeMcients (r2) were O.78 (p<
O.OOI) and O.57 (p<(O.OOI), respectively (Table 2). On the other hand, a highiy
significant positive relationship (r2=O,80,p<O.OOI) was found between rvagitariqpsis
spp. and the 10-20ym fraction.

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Diatom community under the iee insummer 17

Table 1.List of species of diatoms observed at 1m belew the fast

ice in the water column at St. A duringthe periodfrem

27 December, 1997 to 3 February, 1998.

DiatomsCentrales

Actinocyeltts actinochitus Simonsen

(Ehrenberg)
Chaetoceros sp, 1sp.
C,Coscinodiscus 2asteromphatus
Ehrenberg
c,Eucampia sp.antaretica

(Castracttne)
Mangin
pseudodenticulata(Hustedt)
PorosiraRhizosolenia Jouse
sp.mierotrias

Steilarima (Ehrenberg)Hasle & Sims

7:halassiosira australis M. Peraga]lo
T tumida (Janisch)Hasle

Pennales
F7agiiariopsis curta (VanHeurck) Hustedt
(Grunow)Krieger
ENaviculaIV.N.NitzschiaN.Pinnularia
cvlindrus

directa(W.Smith) Ralfs
gtacieiVan Heurck
spp.stetlata

Manguin
spp.quadratarea
var, constricta (Ostrup)
Heiden
Pleurosigma directunlGrunow
turgidutoides <Hasle)
ltsceudo-nitzschia Hasle
Stauroneis spp,

Sinking diatoms
Of the dominant diatoms, the two large species, P. pseudodenticulata and R
turgiduloides, and the smatl species ofjF}'agilariopsis showed an opposite trend in cell flux
at 20m below the ice (Fig.7). The cell fluxes ofthe two large species were high during
the first halfof January,with maximajust beforethe middle of January of 5,1 × i06cells
?day'
m for P. pseudodenticulata(Fig.7A) and 1,5× 10S cells m 2dayi for R
turgiduloides (Fig. 7B). Thereafter, their cell fluxes decreaseddrastically to lessthan
2.5× 10Sand 2.8× 107cells m' ?day ], respectively, at the end ofJanuary. Although the
2day'i
cell fluxof the small species was relatively modest at < 1.8× 10' cells m in the first
halfof January,itincreased laterand reached the maximum flux of 1.2× 109 cells m 2
day'i from the end of January to the beginningof February (Fig. 7C). The cells
deposited in the trap were well aggregated.

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18 A. Ishikawa,N. Washiyama, A. Tanimura and M. Fukuchi

1OO
tw-pazChaetoceros sp.1

Porosira
80 pseuciodenticulata
thalassiosira
aushalis

Others
60

40
.AL8voo=cuvc=Dcuo>'.-"-.Jas-ocr

20

o1OO

-hegilariopsis spp.

Pseucto-nitzschia
80 tuigiduioides
Others
60

40

20

o
27 1 10 20 30 3
Dec. Jan. Feb.
1997 t998

Fig. 6. Daily ehanges in relative abundance of the two dominant species umong (A)contric and (B)
pennate diatom populationsat 1m below the ice in the water column at St.A. ND and NC
denote no data Hnd no cells. respectively.

Table 2.Relationships between cell densitiesof3 dorninunt diatoms and

concentrations of size-fraetionated chlorophyll a (10'20"m and //'20
"m) aL lm betow the ice, Coefficient of rcgression
(r!)in
'F denote positiveand negative
parenthesis. and relationships,

respectively,

Fraction of chlerophyll a
Diatom species
',,
le-20gm 20um

Porosirapseudodenticulata - (O.17)- - (O 78)'

m (O.2s)r
Aeudo-nitzschianirgiduioides (O.57)'
1-'agilariopsis - CO.80).
spp, (O.18)#
'
Significantcorre]ation,p ･,'･'
O.OO1.
# ','O.
Non significant correlation, p 1,

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Diatem community under the ice insummer 19

6(A)

ATvnyEseMo 5

3
ooor

2
xv
1

o
18
16Ar {B)
× 14vc>J
==ooen=y'.!co
12E9
tOgsr".

6V,v

2o14

(c)
12ATv

10cy

E8seig
6co04r5

2o

2 10Jan.20 30 2
Feb.
1998
Fig. 7.Temporal changes in sinking cell fiuxesof dominant diatoms, (A)Ilorosira pseudodenticulata,
(B) Ilseudo-nitzschia
turgiduloides and (C)Fhagilariopsis
spp,, rnonitored by sediment trap 20
rn below the bottorn of the fast ice at St. A from 2 January to 2 February 1998. Bars

represent " SD, Note differentscales ony-axes.

Discussion
In thisstudy, the concentration of >20pam chlorophyll afractionat 1 m below the
icewas related to the cell abundances of two largespecies, R pseudodenticulata and R
turgiduloides 2). A higher
(Table significant positiverelationship was found for the
former species although itscell abundance was much lower than that of the latterone.

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20 A. Ishikawa, N. Washiyama, A. Tanimura and M. Fukuchi

This isprobably due to the largercell volume of the former species, which contains more

chlorophyll a ina cell than the latter

species. This indicates that duringthe firstperiod
of high totalChl a concentration, i.e.,from the end of December to early January,the
two species represented the phytoplankton population and, furthermore, P.
pseudodenticulata was the primary component of the totalCh) a. These two diatoms
commonly occur in the icealgal assemblage in the Antarctic (e.g., Garrison and Buck,

1989; Garrison,1991) and they have also been found in the sea ice around Syowa
Station(Watanabe, 1982, 1988;Watanabe et aL, 1990). Apart from the ice,limited
information(Watanabe,
pers. commun.) about the species composition in the water

column around thatR pseudodenticulata and P, turgiduloidesoccurred

the station shows

as dominant species among the diatom populationfrom spring to early summer at the

end of 1983 and the cell abundance of the latter

species was much higher than that of the
forrnerspecies. His observation issimilar to ours, suggesting the possibilitythat these
species are major components of the diatom population in the water celumn in early
summer the station.
around On the other hand, a highly significant positiverelation-
ship between density of -'agitariopsisspp. and the concentration
cell of the 10-20ptm

chlorophyll a fraction at 1 m below the ice was found (Table 2), indicating that the
main component in the phytoplankton population duringthe other periodof high total
Chl a, i.e., from the middle to the end of January,was the small nano-sized pennate
diatom. Those species were also observed to be dominating in the water column under
the icearound Syowa Stationin January 1984 (Watanabe, pers. commun.).
Iwanami et al. (1986) observed a development of chlorophyll a concentration at a

pycnocline fbrmed at shallower depth (ca. 1.3m below the ice) near Syowa Station,
from the middle to the end of January 1984,and suggested that the developmentwas due
to growth of phytoplanktonat the shallower pycnoclinewhich kept them from sinking
te poorly lighted deeper layers for a longer time. In this study, instead of no
appearance of an obvious pycnocline, a densitygradient developedthrough the water
column by the end of January (Fig.2C).Furthermore, the apparent growth rate
calculated frornthe change in diatom cell density(Fig. 5A), from 2.6× 10`cells l i to
4.7× 106 cells l i
(themaximum value during the study) on 10 and 22 January,
i)
respectively, was much higher (O.43d than the growth rate of the phytoplankton
community January (O.18d i; Ishikawa, unpubiished
on 16 data), as estimated in situ
(incubated at ca. O,5m below the ice) by the dilution method (Landry and Hassett
1982). This indicates that the developmentof the population at 1 m below the iceafter
10 Januarycannot be explained only by the growth of the cells. jF}ragilariopsis curta and
]Fl
cylindrus are common and often dominate in the ice algal assemblage around Syowa
Station (Watanabe,
1988; Watanabe et al., 1990) and in other Antarctic regions (e.g.,
Burkholderand Mandelli,1965;Richardsonand Whitaker,1979;Garrisonet aL, 1986;
Garrisonand Buck, 1989;Garrison,1991). Ithas been reported that the abundance of
chlorophyll in the bottom ice decreases markedly
a in January with ice melt (Sasaki and
Hoshiai, 1986; Watanabe et aL, 1990 ). Judgingfrom the fact that the ice also melted
in January, 1998, it can be strongly a$sumed that a largenumber of the cells of
iF},agilariopsis spp. were released into the water column from the bottom ice,where those
species dominateover other populations ofice algal species. This isa major reason why
the 10-20pam fractionof chlorophylla in terms of F>'agilariopsis spp. populations

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Diatom cemmunity under the ice in surnrner 21

developedinthe water column from the middle to the end of January. In other words,
itislikelythat most of the phytoplanktoncells in the water column were derived from
the ice,while they could stM grow. Sasakiand Hoshiai (1986) reported that the ice
algae detachingfrom the ice increased in lateDecember, also indicating the possibility
that the phytoplankton increasewe observed at the end of December was a result of the
increaseddetachment of icealgae from the ice.
Ithas been reported that the icealgae released from the iceare a seed for the bloom
in the water column at the iceedge zone (Garrison and Buck, l985;Smith and Nelson,
1985;Garrisonet al., 1987). On the other hand, ithas also been suggested that the
aggregated ice algae sink rapidly to the bottom and do not act as a seed for the bloom
(Riebesell et aL,1991). In this study, relatiyely higherconcentration of total Chla
extended from the surface to the subsurface (Fig. 3A) and the sinking cell fluxof the
dominant species (Fig. 7) increased with decreasingchlorophyll a concentration (Figs.
3A and 4A), indicating that our results would be comparable with the latter scheme.
According to Marchantet aL (1996), the organic materials (marine snow) collected by
divers below the surface in the water column around Syowa Stationinmid-January 1994
were dominated by diatoms and muci]age which derivedfrom the ice community.
Sasaki and Hoshiai (1986) also reported that the icealgae detachedfrom the icearound
the station sink directly to the bottom duringsummer and are net grazedduring sinking.
Thus, the algae reaching the bottom become a major fbod source for the benthic
animals.
As a whole, abundance and community structure of diatoms varied drastically
within the short summer of 1997/98.
The variation was due predominantlyto abiotic
events, such as ice rather than bioticevents,
melting, such as growth of the

phytoplankton. The hydrographic conditions during this summer, especially warm

water exceeding OOC and extension of the salinity gradientto the bottom layerin the
water column, might be abnormal when compared to the conditions usuaily reported
around Syowa Station(Wakatsuchi, 1982; Iwanami et aL, 1986; Satoh et al., 1986;
Fukuchi et al.,1984, 1985). Nevertheless, the highest abundance of phytoplankton in
the water column usually has been reported from the end of Januaryto the beginning of
February (Hoshiai, 1969;Fukuehi et aL, 1984; Iwanami et aL, 1986; Satoh et aL,1986),
as observed in this study. Satohand Watanabe (1988) reported an increaseof the
relative importance of the phytoplankton as the primary producers in the summer
season. Although the relationship between both communities, ice and sea water, in the
ice covered area at the north of Syowa Stationhas been examined before(Hoshiai and
Kato, 1961),itisstill equiyocal. Furtherinvestigations on composition and succession
of phytoptankton species in both water and ice in relation to variables in the en-
vironmental conditions during the austral summer are needed to clarify the fateof the
ice algae and the origin of the phytoplankton in the water column, leading to better
understanding of the carbon cycle of the coastal ecosystem associated with ice in the
Antarctic during the austral summer.

Acknowledgments
We are gratefulto Profs, K. Moriwaki and K. Shibuya, the leaders of the 39th

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22 A. Ishikawa, N. Washiyama.A. Tanimura and M. Fukuchi

Japanese AntarcticResearchExpedition, for givingus helpfu1 advice on the field work.

We a}sQ express our thanks to the members of the 39th and 38th parties, especially Mr.
A. Masuyama and Dr. K, Seto, respectively, for their cooperation in the field work.

Thanks are due to the HydorographicDepartment,MaritimeSafetyAgency, for lending

us the CTD through the participation of Mrs. M, Yoritaka(39th party). We appreci-
ate the kindnessof Dr. K. Watanabe for providingus with information about the field
work and for his guide to references. Our special thanks are extended to Dr. Harvey
J. Marchant, Australian Antarctic Division,forhisencouragement and critical reading

of this manuscript.

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rReceived
October 13, l999; Revised manuscript accepted FlebruatyZ 2000)

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