The Holocene 14,2 (2004) pp.

246-257

Coastal oceanographic conditions in the

Prydz Bay region (East Antarctica) during
the Holocene recorded in an isolation
basin
Elie Verleyen,l * Dominic A. Hodgson,' Koen Sabbe,
Koenraad Vanhouttel and Wim Vyverman1
('Lab. Protistology and Aquatic Ecology, Ghent University, Krijgslaan 281-S8,
B-9000 Gent, Belgium; 2British Antarctic Survey, Natural Environment Research
Council, High Cross, Madingley Road, Cambridge CB3 OET, UK)
Received 21 October 2002; revised manuscript accepted 28 January 2003

Abstract: Information on East Antarctic coastal environments during the Holocene is relatively sparse. This
is surprising as sedimentary records from the interface between land and sea can provide chronologies of
climatic change, isostatic uplift, relative sea level and the colonization of newly formed biomes. Here we
examine a sediment core from Pup Lagoon, a coastal lake in the Larsemann Hills, East Antarctica. Sediment
stratigraphy, fossil pigments and diatoms were used to infer the sequence of Holocene environmental and
climatic change. Results show that between 5800 and 5500 cal. yr BP the marine coast of Prydz Bay was
characterized by stratified, open-water conditions during spring and summer and seasonally warm conditions.
From 5500 to 2750 cal. yr BP sea-ice duration in Prydz Bay increased with the coast being ice-free for 2-3
months each year, conditions which are similar to the present day. A return to stratified, open-water conditions
and a reduction in winter sea-ice extent between 2750 and 2200 cal. yr BP is signalled by enhanced biogenic
production and more open-water diatom taxa. This is consistent with evidence for the mid-Holocene Hypsither-
mal detected in other records in East Antarctica. Isostatic isolation of the Pup Lagoon basin from the sea
between 2200 and 2000 cal. yr BP slightly precedes the emergence of lakes with comparable sill heights from
A the nearby Vestfold Hills. The colonization of Pup Lagoon after its isolation as a freshwater lake was initiated
HOLOCENE by a siliceous flora dominated by stomatocysts with microbial mat development being prevented by mechanical
RESEARCH or physical stress. A brief period of marine incursion following the mid-Holocene Hypsithermal may be related
PAPER
to local events such as iceberg calving or to minor sea-level change. Weighted averaging regression, used to
infer salinity in the lacustrine zone, shows that from 1500 cal. yr BP Pup Lagoon is a freshwater lake, where
the flora is dominated by stratified cyanobacterial mats, with green algae and diatoms as co-dominants, compa-
rable to modern Pup Lagoon and other lakes in the Larsemann Hills.

Key words: Isolation basin, coastal oceanography, Larsemann Hills, palaeolimnology, diatoms, stomatocysts,
pigments, ice-sheet retreat, Hypsithermal, relative sea-level change, East Antarctica, Holocene.

Introduction and have enabled reconstructionsof offshore marine environments

Information on near-shore coastal environments of East Antarctica
during the Holocene has largely been restricted to studies of mar-
ine sediment cores collected from the continental shelf areas
beyond the limit of iceberg disturbance (e.g., Taylor et al., 1997;
2001; Domack et al., 1998). These cores have answered questions
on the extent of the ice sheets at the last glacial maximum (LGM)
*Author for coirespondence (e-mail: elie.verleyen@rug.ac.be)
Arnold 2004
(past sea-ice extent and inferred surface water temperature). More
recently, attention has been focused on the near-shore environ-
ment, where sediment cores have revealed biological colonization
and succession in marine environments over glacial-interglacial
cycles and late-Quaternary geomorphological and environmental
change. Sedimentary studies of coastal lake basins, inundated by
the sea prior to isostatic uplift in the Holocene are of particular
interest as they permit the determination of changing coastal mar-
ine environments, the timing of isostatic uplift, relative sea level
10.1 191/0959683604hI702rp

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 Elie Verleyen et al.: Coastal oceanographic conditions, East Antarctica, from an isolation basin 247
and the colonization of newly formed biomes. In particular, low-
altitude coastal lakes can permit an accurate reconstruction of the
history of marine transgressions. This is done by dating the tran-
sition between marine and lacustrine assemblages, which has pre-
viously been used to determine the relationship between relative
sea-level change, ice thickness and isostatic rebound in East Ant-
arctica (Zwartz et al., 1998).
The specific aims of this study were to document the history
of environmental change using a sediment core from Pup Lagoon,
a coastal lake on Stornes in the Larsemann Hills, East Antarctica
(Figure 1). Multiproxy analysis was carried out using sedimen-
tology, diatoms, other microfossils and fossil pigments. In combi-
nation, this approach is used to (1) determine main periods of
environmental change in the marine environment, (2) date the
transition between marine and freshwater assemblages and com-
pare these results with the relative sea-level history in the Vestfold
Hills (Zwartz etal., 1998) and (3) reconstruct the succession of
autotrophic biota and environmental change in the newly isolated
freshwater lake.
To enable the interpretationof the fossil data, data sets compris-
ing information on lake-water chemistry, community structure and
composition of modern microbial mats from more than 50 refer-
ence lakes were used to infer environmental conditions at the time
of deposition (Hodgson et al., 2001b). For diatoms, quantitative
inferences of lake conductivity were made using a recently
developed diatom-conductivity transfer function with weighted
averaging calibration based on the East Antarctic calibration data
set (Verleyen et al., 2003).
Site description
The Larsemann Hills (69°23' S, 76°53' E), Prydz Bay, is an ice-
free area on the Ingrid Christensen Coast, Princess Elizabeth
Land, located approximately midway between the eastern
extremity of the Amery Ice Shelf and the southern boundary of
the Vestfold Hills. The region consists of two main peninsulas,
the western named Stornes and the eastern Broknes/Mirror,
together with a number of scattered offshore islands (Figure 1).
An extensive description of the geology, physiography and cli-
mate of the region is given in Stuwe et al. (1989), Gillieson et al.
(1990) and Hodgson etal. (2001a). At 50 km2 the Larsemann
Hills constitute the second largest of only four major ice-free
0 kilometres
Figure 1 Map of the Larsemann Hills, showing the location of Pup Lagoon (L23).
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0 lwmwm.
oases containing lakes along East Antarctica's 5000 km of coast-
line (Fogg, 1998). Recent data (Hodgson etal., 2001a) have
shown that many lakes in this region have accumulated long sedi-
mentary records with radiocarbon dating showing consistent age-
depth sequences. Consequently, these lakes are of great potential
interest for palaeoenvironmental and palaeoclimatological studies,
complementing and extending information obtained from lakes in
the Vestfold Hills (e.g., Roberts and McMinn, 1998) and the
Bunger Hills (Roberts et al., 2000).
Pup Lagoon is a shallow freshwater lake situated 100 m from
the coast of the Stornes Peninsula. Thirty-four percent of the
peninsula is covered by multiyear snowfields (Gore, 1997). The
lake is freshwater, about 1 ha in size with a maximum depth of
6 m and with a sill height of 4 m above sea level (Gillieson et al.,
1990). Pup Lagoon was first studied by Gillieson (1991), whose
short sediment core indicated that biogenic sediment had
accumulated and that microfossils were well preserved.
Methods
During November 1997, a 3.02 m sediment core was extracted
from the deepest point of Pup Lagoon using a combination of a
Glew gravity corer (Glew, 1991) for the surface sediments and
a square rod operated Livingston corer for intermediate to
basal sediments. The core was photographed, macroscopically
described, sectioned in the field and frozen until analysis. Analy-
ses of sediment wet density, dry weight, organic and carbonate
content were carried out in the laboratory using standard methods
(Dean, 1974).
Radiocarbon dating
Eight samples from the Pup Lagoon core were dated using AMS
'4C by the UK Natural Environment Research Council Radio-
carbon Laboratory. Where possible, samples were derived from
discrete biological remains of cyanobacterial mats and their
associated algal communities. The results are reported here as
conventional radiocarbon years BP (relative to AD 1950) and cali-
brated using the program CALIB 4.3 (Stuiver and Reimer, 1993;
2000). Details on radiocarbon dating of the Vestfold Hills' cores
can be found in Zwartz et al. (1998) and references therein.
10 Projection: Lambert confocal conic
248 The Holocene 14 (2004)
Microfossils
Subsamples for diatom and stomatocyst analysis were taken at 1
cm intervals in the upper 20 cm of the core and at 5 cm intervals
from 20 to 302 cm. The organic matter in the samples was
digested with H202 (30 %) and CH3COOH (95%) following
Renberg (1990). A solution of microspheres with known concen-
tration was added to permit absolute diatom counts (Battarbee and
Kneen, 1982). Naphrax was used as mounting medium. At least
400 valves (>2/3 intact) and/or stomatocysts were counted in
each sample.
Pigments
Benthic pigments were extracted from bulk sediments following
HPLC analysis protocols described in Hodgson et al. (1997; 1998)
and Leavitt and Hodgson (2001). The HPLC components were a
Kromasystem 2000 HPLC with a Kontron pump, auto sampler
and diode array detector. The column used was a 25 cm X 4.6
mm reverse phase Spherisorb ODS-2 with a particle size of 5 m.
Further developments in this method have been recently reported
in Airs etal. (2001). The system was calibrated to reference
cultures following SCOR protocols (Jeffrey et al., 1997) and US
Environmental Protection Agency Standards. Total benthic
chlorophylls and carotenoids are expressed as organic matter spe-
cific concentration (ng/g TOC), because comparison of long-term
live plankton data with varved fossil records indicates that this
metric most accurately captures variations in algal abundance
(Leavitt and Findlay, 1994; Leavitt etal., 1997; Verschuren et al.,
1999). The abundance of individual pigments is presented as a
percentage of total chlorophylls or carotenoids in order to exclude
the influence of the organic matter specific concentration.
Ecological modelling and statistical analyses
Statistical exploration of patterns in the diatom and pigment strati-
graphies was performed using indirect ordination analysis. All
data were logarithmically transformed in order to reduce skewness
and to obtain normality in their distributions. Multivariate analy-
ses were carried out using Canoco 4.0 for Windows (ter Braak
and Smilauer, 1998). Fossil diatom data were interpreted using a
reference data set of modern diatom assemblages and environmen-
tal data from 56 lakes from the Larsemann Hills and the Bolingen
Islands. Stratigraphic data were zoned using CONISS, a strati-
graphically constrained cluster analysis following squared root
transformation and plotted using TILIA 2.0b4 (Grimm, 1991),
TILIA*GRAPH 2.20 and TGView version 1.1.1 .1 (Grimm, 2001).
Only taxa found at more than 2% abundance were incorporated
in the CONISS (Grimm, 1987) analysis. Palaeosalinity was
inferred using CALIBRATE 1.01 (Juggins and ter Braak 1997-
2001) with the East Antarctic diatom based weighted averaging
salinity transfer function (Verleyen et al., 2003).
Relative sea-level history
During a sea-level highstand, marine basins close to the shore
accumulate marine sediments. As sea level falls, these basins are
isolated and, if they have freshwater input, are flushed and start to
accumulate freshwater lacustrine sediments. By dating the marine-
lacustrine transition in sediment cores from these lake basins and
by measuring the elevation of the former threshold (sill height),
a precise sea-level record can be obtained. The sill heights in the
Vestfold Hills were measured using a theodolite and electronic
distance meter by the Australian Division of Mapping with an
accuracy of 5 cm. In the Larsemann Hills, sill heights are derived
from spot heights on maps, with an accuracy of 0.5 m. The
obtained sea-level record can be related to ice thickness and iso-
static rebound (e.g., Svendsen and Mangerud, 1987; Zwartz et al.,
1998). The main advantage of this method is that no marine reser-
voir correction is needed if the first freshwater sediments are radio-
carbon dated. In contrast to the previously published sea-level
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record from the Vestfold Hills (Zwartz et al., 1998), radiocarbon
dates are corrected for the marine reservoir effect and calibrated.
Results
Lithostratigraphic analysis
The uppermost 145 cm of the Pup Lagoon core is an organic-rich
sediment consisting of green/brown cyanobacterial mats with high
loss-on-ignition (%LOI) values (Figure 2). From 145 to 150 cm
the microbial mats are decayed. From 150 cm to 302 cm the
organic content rapidly declines with generally less than 10%
organic content, with a peak between 185 and 200 cm. Between
250 and 255 cm sandy layers with gravel are found, associated
with a peak in dry weight (%D.W.) and a minimum in %LOI.
Below 300 cm the core consists of compressed sediments, associa-
ted with an increase in %D.W.
Radiometric dating and sedimentation rates
The surface date (Table 1) reveals that 'IC in modern samples is
near equilibrium with atmospheric `4C; consequently, a correction
for the 'reservoir effect' in freshwater sediments is not needed
(Hodgson etal., 2001a). The AMS 14C dates of the marine sedi-
ments are corrected for the 'reservoir effect' by subtracting 1300
years from the original '4C date, following recent conventions for
the Antarctic marine environment (e.g., Berkman etal., 1998;
Ing6lfsson et al., 1998). The sedimentation rate is relatively high
(0.1763 cm yr-') between the base of the core and 250 cm and
decreases rapidly until 200 cm (Figure 2). Between 200 and 50
cm the sedimentation rate is relatively stable and high, after which
it decreases gradually.
Pigment analysis
The pigments recovered from Pup Lagoon and their taxonomic
affinities are listed in Table 2. Total carotenoids and total
chlorophylls are expressed as organic matter specific concen-
tration (ng/g TOC). Each separate pigment is expressed as the
abundances (in %) of the class to which it belongs (Figure 3).
Constrained cluster analysis in CONISS (Grimm, 1991) suggests
that the core can be divided into two distinct zones: PLPAI (Pup
Lagoon Pigment Analysis 1) and PLPA2. PLPAI (302-140 cm,
c. 5800-2000 cal. yr BP) is characterized by a relatively high
amount of total carotenoids and chlorophylls. Two bacteriophaeo-
phytin derivatives and one bacteriochlorophyll dominate the
chlorophylls in PLPAI and diatoxanthin and dinoxanthin are the
most abundant carotenoids throughout this zone. PLPA1 is further
divided into three subzones: PLPA1A, PLPA1B and PLPA1C.
The uppermost layers of PLPA1C (between 155 and 140 cm,
c. 2200-2000 cal. yr BP) are characterized by a mixture of pig-
ments, specific for PLPA1 or for PLPA2. Total carotenoids and
total chlorophylls are significantly lower in PLPA2 (140-0 cm, c.
2000 cal. yr BP to present) with the exception of a single sample
at 18 cm. Lutein, zeaxanthin, nostoxanthin and echinenone are
the dominant carotenoids, and bacteriochlorophyll 2 derivative,
phaeophorbide a and chlorophyll a are the most abundant
chlorophylls.
Microfossil analysis
Seventy-eight samples were analysed and 115 diatom taxa were
identified. Twenty-nine taxa are found at more than 2 % abun-
dance. A list of the lacustrine diatoms of the Larsemann Hills and
their taxonomic authorities is given in Sabbe et al. (2003). For
marine species the reader is referred to Roberts and McMinn
(1999). Three distinct diatom zones are recognized on the basis
of the ordinations and the CONISS analysis, PLDAI (Pup Lagoon
Diatom Analysis 1) PLDA2 and PLDA3 (Figures 4, 5 and 6).
PLDA1 (302-150 cm, c. 5800-2140 cal. yr BP), is dominated
 Elie Verleyen et a!.: Coastal oceanographic conditions, East Antarctica, from an isolation basin 249

Uthology age
0 1000 2000 3000 4000 5000 6000 7000 0 20 40 60 80 100
0 I,4- - o-
14C dates (yr BP)
50 -
- - - calibrated dates (yr BP) 50 -

100 - 100 -

E 150 - 150 -

s al
-S
.X 200 - 200 -

-~~~~~~~~~~~~~~~'
250 250 - P

300 - 300 - 4

350 - - 350 -
microbial mats -
crudely stratified
microbial mats compressed/decayed
-

consolidated organic sediment

consolidated sediments sands and grits

Figure 2 Lithology, lithostratigraphic analyses (%LOI, %D.W.) and age-depth curve for the Pup Lagoon core.

Table 1 Radiocarbon age, calibrated ages and publication codes core levels (155-0 cm) with the modern flora of the Larsemann
Hills (Sabbe et al., 2003) a DCA was run with the fossil samples
Depth Conventional Publication Corrected Calibrated included as passive samples (Figure 6). This information can help
(cm) lC age code 14C age age to infer the environmental conditions prevailing during deposition
(yr BP ± lr) (yr BP) (cal. yr BP) of fossil assemblages. Two distinct clusters can be recognized in
the surface sample set described in Sabbe etal. (2003). Lakes
0 Modem AA-35718 - 122 deeper than 2.5 m exhibit a different flora compared to the shallow
20 675 ± 40 AA-35748 - 653 ponds (<2.5 m), where well-developed benthic microbial mats
50 1270 ± 40 AA-35749 - 1204 are absent. This is possibly due to mechanical and/or physical
100 1590 ± 45 AA-35750 - 1517
150 2150 ± 45 AA-35751 -
stresses, resulting from ice freezing to the bottom for most of the
2139
200 3915 ± 45 AA-35752 2615 ± 45 2749 year. Samples from PLDA2 and the sample at 155 cm depth are
250 6085 ± 50 AA-35753 4785 ± 50 5504 similar to the shallow ponds, where stomatocysts dominate the
300-302 6380 ± 50 CAMS-50377 5080 ± 50 5794 siliceous protist community. PL 135-110 are clustered near lakes
which are more than 2.5 m deep and with a conductivity of more
than 2 mS/cm. PL 105-0 are clustered around the modern diatom
assemblage of Pup Lagoon and are similar to the deeper lakes
by marine and sea-ice associated diatoms (e.g., Fragilariopsis (>2.5 m) with a conductivity of less than 2 mS/cm. In the fresh-
cylindrus, Fragilariopsis curta and Navicula glacie). The number water zone, salinity can be inferred using a diatom-based transfer
of frustules per gram sediment dry weight is significantly higher function for East Antarctica. Jack-knifed r2 and apparent r2 are
in this zone compared with the rest the core. PLDA1 is subdivided high in the East Antarctic transfer function, 0.83 and 0.86 respect-
into three zones: PLDA1A, PLDA1B and PLDA1C. These closely ively and RMSE (0.27) and RMSEP (0.31) are relatively low.
correspond to the zones identified in the pigment analysis. The reconstructed salinity is relatively higher and more variable
Chaetoceros resting spores are relatively abundant in PLDA1A, between 160 and 95 cm, compared with the upper core levels
at 225 cm and in the basal levels of PLDA1C. The upper layers
(Figure 7). The highest modelled salinity occurs at 160 cm.
of the latter zone are dominated by the brackish-water diatoms
Navicula phyllepta and Craspedostauros laevissimus. PLDA2 Relative sea-level change
(150-140 cm, c. 2140-2000 cal. yr BP). is dominated by stomato- The relative sea-level history of the region is obtained by plotting
cysts, presumably siliceous resting stages of Chrysophyceae and the ages of lacustrine-marine and marine-lacustrine transitions in
by lacustrine brackish-water diatoms. The diatom assemblage in the lake cores against the sill height of the lake. The curve passes
PLDA3 (140-0 cm, c. 2000 cal. yr BP to present) is dominated through these points or the error bars plotted to allow for the
by the freshwater diatom Stauroforma inermis and by Pinnularia uncertainty in measurements of sill height and error ranges of
microstauron, The euryhaline diatom Amphora veneta is more radiocarbon dating. For the Vestfold Hills lakes, dates are extra-
abundant in the lower levels of this zone (PLDA3A, 140-95 cm, polated, assuming a constant sedimentation rate. Furthermore,
c. 2000-1500 cal. yr BP). In a single sample at 100 cm depth dates were corrected for the reservoir effect by subtracting 950
Planothidium quadripunctatum occurs, together with freshwater and 1310 years (Zwartz etal., 1998, and references therein).
(Psammothidium abundans), and brackish-water (Craspedostauros These uncertainties are included in Figure 8. No corrections were
laevissimus) diatoms and some marine and sea-ice associated applied to the date of isolation of Pup Lagoon, since lacustrine
diatoms (F. curta). sediments were dated. An additional source of uncertainty is the
In order to compare the lacustrine diatom assemblages of the exact timing of the isolation of the lake. Maximum peak-to-peak

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250 The Holocene 14 (2004)

Table 2 Pigments commonly recovered from Pup Lagoon sediments and their taxonomic affinities. Quantitatively important sources indicated in italics.
Local interpretation based on Jeffrey et al. (1997). Predominant source identified as planktonic (P), littoral (L) or sedimentary (S) (postdepositional
derivatives). Relative degree of chemical stability and preservation is ranked from most (1) to least (4) stable based on mesocosm experiments and mass-
balance studies (see Leavitt, 1993); - = uncertain stability

Pigment Source' Stability Affinity Local interpretation

Total chlorophylls Relative production

Chlorophyll a P, L 3 all photosynthetic algae Relative production
Chlorophyll b P, L 2 Chlorophyta, Euglenophyta Open-water blooms of green algae
Pheophytin a P, L, s 1 Chl a derivative General algal detritus
Pheophorbide a P, L, s 3 Chl a derivative Grazing, senescent diatoms, general algal
detritus
Pyro-pheo(pigments) L, S derivatives of a and b-phorbins Zooplankton faecal pellets
Chlorin Chl a derivative
Pupurin Chl a derivative
Bacteriochlorophylls S Anaerobic bacteria Sediment anoxia due to ice cover

Total carotenoids Relative production

Alloxanthin p Cryptophyta
Lutein P, L Chlorophyta, red seaweeds Open-water blooms of green algae
Prasinoxanthin p Prasinophytes Seasonal open-water blooms of
Pyramimonas
Fucoxanthin P,L 2 Bacillariophyta, Prymnesiophyta, brown Sea-ice diatoms or open-water diatoms
seaweeds, raphidophytes, some Dinophyta2 (depending on taxa present and mixed
with endosymbionts, Chrysophyta plankton blooms if diadinox. present.
Freshwater diatoms
Violaxanthin L 4 Chlorophyta, Euglenophyta, brown seaweeds Green algal and mixed open-water blooms
(possibly seaweed)
Neoxanthin L 4 Chlorophyta, Euglenophyta, brown seaweeds Green algal and mixed open-water blooms
(possibly seaweed)
Diatoxanthin P, L, s3 2 Bacillariophyta, Dinophyta, Chrysophyta
Diadinoxanthin P, L, s3 3 Bacillariophyta, prymnesiophytes, Seasonal open-water blooms
Chrysophyta, Dinophyta
Dinoxanthin p Dinophyta Seasonal open-water dinoflagellate blooms
Echinenone P,l 1 Cyanobacteria Freshwater cyanobacteria
Zeaxanthin P,l 1 Cyanobacteria, chlorophyta Freshwater cyanobacteria and green algae
Nostoxanthin4 P,L Cyanobacteria Freshwater cyanobacteria
Canthaxanthin P,l 1 colonial Cyanobacteria, herbivore tissues Layered cyanobacterial mats
Myxoxanthophyll P,l 2 colonial Cyanobacteria Layered cyanobacterial mats
Scytonemin5 P,L colonial Cyanobacteria Layered cyanobacterial mats under UV
'stress'
R,P-carotene P, L Chlorophyta, chromophyte algae, some Relative production
phototrophic bacteria
P,S,-carotene P, L 2 Chlorophyta, cryptomonads, red algae Green algae
R,By-carotene P, L Chlorophyta, chromophyte algae Green algae
'Importance of given source also depends on site morphometry and physical properties (e.g., clarity).
'Some dinoflagellates contain peridinin as their main carotenoid, but this very labile compound is rarely recovered from marine sediments.
3Diatoxanthin interconverts with diadinoxanthin in marine sediments. Both are elements of the xanthophyll cycle in chromophyte algae.
'Nostocales and allied cyanobacteria have a highly diverse complement of similar, glycosidic carotenoids. Many of these pigments are not easily separated
by HPLC (cf. Leavitt and Findlay, 1994).
5Cyanobacteria, particularly benthic colonial forms, may produce and deposit scytonemin and its derivatives in some lakes which are transparent to UV
radiation (Leavitt et al., 1997).

tidal range is 1.7 m, implying the connection of the lake to the
sea could lie within this tidal range for more than 1000 years
(Zwartz et al., 1998). The isolation of Pup Lagoon (sill height 4
m) from the sea is complete approximately 1000 years later than
the emergence of lakes with comparable sill heights in the
Vestfold Hills (e.g., Organic Lake and Watts Lake, Figure 8). The
Holocene sea-level highstand in the Vestfold Hills is situated
around 7200 cal. yr BP. The rate of sea-level change ranges
between 2 (Larsemann Hills) and 1.5 mm yr- I (Vestfold Hills).
Discussion
This study describes the first long sedimentary record of environ-
mental change from a sedimentary basin in the Larsemann Hills.
Gillieson (1991) reported preliminary results from a 40 cm deep
core and suggested that Pup Lagoon is a proglacial lake, which
originated after retreat of the ice sheet. The core in the present
study is 302 cm long and the stratigraphic consistency in the age-
depth sequences and undisturbed laminae throughoutthe core pro-
vide new evidence that this lake was a marine basin isolated in
the late Holocene following isostatic uplift. In addition, the
sediments have remained undisturbed by glacially or biologically
mediated remixing. A reliable chronology could therefore be
obtained for the Pup Lagoon core by dating eight depths, includ-
ing the marine-lacustrine transition. Both cluster analysis and
ordination of diatom and pigment data reveal congruent zoning
of these proxies in the core. Three distinct zones can be recog-
nized on the basis of the lithostratigraphic, pigment and diatom
analyses: a shift from marine to lacustrine conditions with a
well-defined transition zone in between. Significant changes in
environmental conditions can be inferred from the different

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 Elie Verleyen et al.: Coastal oceanographic conditions, East Antarctica, from an isolation basin 251

Figure 3 TILIA graph of fossil pigment data. Zoning is based on a CONISS constrained cluster analysis. Each separate pigment is expressed as a relative
abundance (in %) of total chlorophylls and carotenoids. Scytonemin belongs to neither group but is plotted relative to carotenoids for convenience. For
a key to the lithology, see Figure 2.

m1b
-t.
e,. A"AI.Ie !4e-
e; +,
N , O'e
;f ..y
db
4F .

- ."e
Zone

Figure 4 TILIA graph of diatom data with a CONISS constrained cluster analysis. Data are expressed as relative abundances. For a key to the lithology,
see Figure 2.

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Aft
252 The Holocenq 14 (2004)

A related to high productivity (e.g., Leventer, 1992; Leventer et al.,

1993; Barcena et al., 1998) and stratified, open-water conditions
during summer and spring (Whitehead et al., 2001). The high rela-
tive abundance of diatoxantin together with smaller abundances
of dinoxanthin and other carotenoids indicate a diverse and pro-
longed open-water summer plankton bloom. Low concentrations
of bacteriochlorophyll,relative to the rest of zone PLPAI, suggest
that sediment anoxia and occupation by anaerobic bacteria was
not widespread. The sedimentation rate is significantly higher dur-
co ing this period, compared with the rest of the core. No significant
decline in %LOI can be distinguished between this zone and the
rest of zone PL(P)DA1, suggesting that the higher sedimentation
rate is not caused only by inorganic sediment input. This increase
in open water may be due to seasonally warmer conditions, or to
increased wind stress as suggested by Cunningham and Leventer
II (1998). In our opinion, the increase in open water is related to
changed oceanic conditions in this part of Prydz Bay. The acceler-
ated deglaciation of the coastal zone of Prydz Bay during this
period (including Stornes) may have resulted in increased seasonal
meltwater input which could have led to stratified and more open-
water conditions. During this period, maximum colonization by
snow petrels is reported in the Bunger Hills (Figure 10) and linked
B with fast ice retreat (Verkulich and Hiller, 1994). Zhang (1992)
suggests that a climate optimum may have existed in the nearby
Vestfold Hills between 6200 and 3700 yr BP. High abundances
of Chaetoceros resting spores and vegetative cells are also found
in Anderson Lake (Vestfold Hills) prior to 6730 uncorrected yr
BP (Roberts and McMinn, 1998), possibly related to the continu-
II ous deglaciation of parts of the Vestfold Hills during this period
(Pickard et al., 1986). This rapid ice retreat is probably related to
an early-Holocene warm period in Antarctica as reported from
different East Antarctic ice cores (Masson et al., 2000) and from
marine cores (e.g., Cunningham et al., 1999; Taylor et al., 2001).
o
N
Sea-ice conditions, similar to present situation
(c. 5500-2750 cal. yr BP)
Different proxies suggest that the conditions are similar to the
present situation, with Prydz Bay partly ice-free for 2-3 months
each year (Rathburn et al., 1997). The sediment-accumulationrate
(0.0235 cm yr- ') is significantly lower than in the previous zone
and is similar to Holocene sedimentation rates determined from
the Prydz Bay region (Taylor and McMinn, 2002). The sea-ice
associated taxa Fragilariopsis cylindrus, F. curta (Zielinski and
Gersonde, 1997; Barcena et al., 1998) and Navicula glaciei
-1.0 A,=0.23 +1.0 (Roberts and McMinn, 1999), characteristic of the continental
shelf assemblage (Taylor et al., 1997), dominate the siliceous
Figure 5 (A) Principal components analysis (PCA) of the diatoms in the flora. The carotenoid data in this zone show an increase in the
entire core. (B) PCA of the diatoms in the marine core levels (between
302 and 155 cm). The sample at 225 cm depth is clustered in PLDA1A
relative proportion of dinoflagellates (dinoxanthin) and chloro-
because of the high abundance of Chaetoceros resting spores. Axes 1 and phytes (violaxanthin) indicative of a single seasonal open-water
2 are shown. bloom. Chlorophylls (consisting of a bacteriochlorophyll and two
bacteriophaeophytin derivatives) are found in higher concen-
trations in this zone than in any other part of the core. These
proxies within the three main zones (Figure 9). These yield infor- bacteriochlorophylls are characteristic of a high degree of anaer-
mation on coastal oceanographic environmental changes in Prydz obic bacterial activity in the seasonally anoxic sediments that
Bay between 5800 and 2200 cal. yr BP, on coastal limnological would be expected to develop under a prolonged period of annual
changes between 2000 yr BP and present, as well as on the timing ice cover. The presence of Chaetoceros resting spores at 225 cm
of the marine-lacustrine transition providing information on (c. 3900 cal. yr BP if sedimentation rate is assumed to be constant
relative sea-level history. between 250 and 200 cm) could be due to an increase in meltwater
input, corresponding with the ice-sheet retreat in the northeastern
Coastal oceanographic conditions in Prydz Bay part of Stornes and the related start of biogenic sedimentation in
(c. 5800-2200 cal. yr BP) Lake Burgess at c. 4180 cal. yr BP (Hodgson etal., 2001a).

Stratified, open-water conditions (c. 5800-5500 cal.
BP)
The different proxies suggest stratified and open-water conditions
between c. 5800 and 5500 cal. yr BP. Chaetoceros resting spores
are relatively abundant in this zone of the core. These taxa are
yr Stratified, open-water conditions (c. 2750-2200 cal. yr
BP)
Enhanced biogenic production and more open-water diatom taxa
in this period may be related to rapid ice retreat, stratification, a
reduction in winter sea-ice extent and a longer period of

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 Elie Verleyen et al.: Coastal oceanographic conditions, East Antarctica, from an isolation basin 253

3.5

25
co
CS
2
0
0
1.5

3.5 no 0 - '

0 0.5 1 1.5 2 2.5 3 3.5 4 4.5

DCA1 Ak=0.65
Figure 6 Detrended correspondence analysis (DCA) of the Larsemann Hills surface data set (Sabbe et al., 2003) with the freshwater core levels included
as supplementary sites. Samples with more than 10% marine species are not incorporated in the analysis. Axes 1 and 2 are shown.

open-water conditions. A decline in the concentrations of bacteri-

ochlorophylls indicates that sediment anoxia and sea-ice cover
was reduced. Similarly, lutein gradually increases from 210 cm,
and peaks at 185 cm, which may be a signal that rhodophyte algae
were becoming established at the site. These have been reported
from coastal locations in continental Antarctica (Amsler, 1995;
Hodgson et al., 2004). There is also a distinct increase in one of
the diadinoxanthin derivatives, which suggests that there were
well-developed seasonal open-water blooms with representatives
from the diatoms, prymnesiophytes,chrysophytes and dinophytes.
0 20 40 60 80 100
cor depth (e)
120 140 160 180 Brackish-water diatoms increase in abundance towards the top of
this zone. These observations may indicate that the Pup Lagoon
Figure 7 Inferred salinity based on the transfer function for East Antarctic basin and the coastal areas at similar elevations on the Stornes
diatoms (Verleyen et al., 2003) in the lacustrine assemblages of the core.
Salinity is logarithmically transformed.
Peninsula were exposed in the near-shore environment (as dis-
cussed in detail below).

10

9 i. s -
8 - fl
7

E 6 hHh

._ 4 r

0
9000 8000 7000 6000 5000 4000 3000 2000 1000 0

calibrated dates (yr BP)

Figure 8 Relative sea-level history of the Vestfold Hills based on lake isolation observations (Zwartz et al., 1998) with Pup Lagoon (this study) added
to the graph. Triangles under the curve represent terrestrial sediment deposits containing marine assemblages. The dashed line shows the sea-level obser-
vations from the Vestfold Hills. The full line shows the sea-level record in the Larsemann Hills based on Pup Lagoon. 1 = Ace Lake; 2 = Anderson Lake;
3 = Druzhby Lake; 4 Highway Lake; 5 Watts Lake; 6 Pup Lagoon; 7 Organic Lake.
= = = =

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254 The Holocene 14 (2004)

5800- 5500 5500 -2750 2750- 2200 2200-2000 2000 -1500 1500 - 0
caLyr BP caLyr BP caLyr BP cal.yr BP caLyr BP cal.yr BP

marine- marine marine- marine- lacustrine - lacustrine -

ssonal sea ice sasonal freashwater - salinity stabl, sanity
open water open water transition

.- . .---

isostatic
rebound
I brief marine
incursion
(c. 1500 calyr

BP)
Global
sea level
~JfJ Sea ice
Bedrock

Figure 9 Cartoon showing climate and environmental history of the Pup Lagoon basin.

Figure 10 Comparison of Holocene palaeoclimates. References: (1) this study and Hodgson et al. (2001a; 2001b); (2) Roberts and McMinn (1998); (3)
Roberts et al. (1999); (4) Zhang (1992); (5) Roberts et al. (2000); (6) Verkulich and Hiller (1994); (7) Cremer et al. (2001); (8) Masson et al. (2000); (9)
Rathbum etal. (1997); (10) Taylor and McMinn (2002); (11) Bjorck et al. (1993); (12) Jones et al. (2000); (13) Taylor et al. (2001). Dates used in this
figure are as cited in the original source and have not been independently corrected nor calibrated. Dates in ice-core records
(*) are in calendar years BP.

The near-shore conditions reported in this zone are consistent
with evidence for the mid-Holocene Hypsithermal (see Figure 10
for an overview), which has also been detected in other records
in continental Antarctica (e.g., Roberts and McMinn, 1998;
Roberts et al., 2000) and the Antarctic Peninsula (e.g., Bjork
et al., 1991; 1993; 1996; Jones et al., 2000).
Transition between marine and lacustrine sediments
and relative sea-level change (c. 2200-2000 yr cal.
BP)
The lower core levels of zone PLDA2 are characterized by a high
abundance of brackish-water diatoms. The diatom assemblage in
these core levels is very similar to those of modern mesosaline
lakes in the Larsemann Hills. This suggests that at this time the
Pup Lagoon basin was isolated from the sea due to isostatic uplift,
in response to the regional deglaciation. The emergence of Pup
Lagoon can be dated between 2200 and 2000 cal. yr BP, when
recurrent limited transgressions prevented the formation of
microbial mats. This resulted in a weakened persistence of the
marine chlorophytes and euglenophytes and possibly seaweeds
(neoxanthin, violaxanthin). During this period the marine inlet
became brackish, due to meltwater input. Between 150 and 140
cm depth stomatocysts are very abundant. Today, shallow ponds
in the Larsemann Hills are characterized by a high abundance of
these resting stages, whereas well-developed benthic microbial
mats are absent, probably as a result of the mechanical and/or
chemical stress imposed by ice freezing to the bottom. We suggest
that chrysophytes are among the first to colonize newly exposed
Antarctic isolation lakes, in which ice-cover dynamics or other
forms of physical stress (e.g., limited tidal influence) prevent the
formation of cyanobacteria-dominaed microbial mats. Despite
this, non-mat-forming cyanobacteria were present in this zone,

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 Elie Verleyen et al.: Coastal oceanographic conditions, East Antarctica, from an isolation basin 255
and in the lower parts were under considerable UV stress (a peak
in the UV screening pigment scytonemin at 150 cm). Similar pig-
ment compositions are found today in some of the shallowest
lakes in the Larsemann Hills where the cyanobacteria mats occur
as small flakes and biofilms. The establishment of the diatoms
and green algae towards the top of the zone is reflected in the
increase in fucoxanthin, neoxanthin and violaxanthin.
The timing of isolation of Pup Lagoon (Figure 8) precedes the
emergence of lakes in the Vestfold Hills with comparable sill
heights (Zwartz etal., 1998), although the age-elevation curve
passes through the error ranges of the Pup Lagoon core. This may
be due to inaccurate measurement of the sill height of Pup Lagoon
or to including the data derived from Laternula species. To date
this bivalve is known to live deeper than I m (Dell, 1990; Ahn,
1994), whereas the sea-level record from the Vestfold Hills
(Zwartz et al., 1998) implies they grew very close to sea-level.
If this 1 m ecological threshold is adapted, sea-level lowering is
approximately similar in both regions. Alternatively, the lakes in
the Larsemann Hills indeed emerged later than the lakes in the
Vestfold Hills with comparable sill height due to a higher local
ice loading and associated crustal depression. This is not unlikely,
since Huybrechts (2002) modelled the crustal depression of the
Antarctic continent and suggested that the region of the Lambert
Glacier-Amery Ice Shelf is a key area in the history of the East
Antarctic Ice Sheet (EAIS). The model indicated a crustal
depression in the Larsemann Hills area between 100 and 200 m
and in the Vestfold Hills between 100 and 50 m around 16000
cal. yr BP. These differences in crustal depression are likely to
affect the late-Holocene rate of isostatic uplift. Furthermore, the
calculated rate from the Larsemann Hills agrees well with rates
inferred by using two out of three ice-sheet models and associated
melting scenarios (c. 1-3.5 mm yr- 1; Zwartz et al., 1999), but it
is lower than the rate calculated in the Windmill Islands (c. 5 mm
yr- ; Goodwin and Zweck, 2000). Furthermore, the Holocene sea-
level highstand c. 7100 years at c. 9 m in the Vestfold Hills is,
compared with other East Antarctic regions, different in altitude
but comparable in age (e.g., S6ya Coast (Miura et al., 1998): 17 m
around 7663 cal. yr BP, Windmill Islands (Goodwin and Zweck,
2000): 31.5 m around 7676 cal. yr BP). The analysis of cores
from other isolation lakes in the region will enable us to calculate
the rate of isostatic uplift and Holocene marine limit more accu-
rately.
The biotic development of a coastal lake and a brief
marine incursion (c. 2000 cal. yr BP to present)
After isolation, laminated microbial mats are well developed and
dominate biological activity in the lake. The pigment composition
indicates dominance by cyanobacteria (scytonemin, nostoxanthin,
zeaxanthin and echinone), with diatoms (fucoxanthin) and green
algae (lutein, ,B,-carotene) as co-dominants. Chlorophyll a,
bacteriochlorophylls and a series of degradation products domi-
nate the chlorophyll signal in this zone. The higher abundance of
the mesohaline diatom Amphora veneta and the relatively higher
and fluctuating inferred salinity could reflect the slow and inter-
mittent input of freshwater into the basin. The relatively slow
dilution of the lake water may be linked to the drier period in the
Bunger Hills suggested by Roberts et al. (2000) on the basis of
an increase in palaeosalinity from 1900 yr BP until present. At
100 cm depth a limited input of marine diatoms is seen, together
with diatoms typical for slightly deeper lakes (Psammothidium
abundans and Planothidium quadripunctatum) as discussed in
Sabbe et al. (2003). A. veneta is abundant at 95 cm and inferred
salinity is higher. These data suggest that a limited marine water
input, due to storms, iceberg calving and the creation of swells
or a small ice readvance, caused a minor increase in salinity and
water depth. After 95 cm (c. 1500 cal. yr BP) the reconstructed
salinity remains relatively stable and the fossil assemblages in the
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core levels are comparable to the surface sample of modern Pup
Lagoon (Figures 6 and 7).
An alternative explanation for this brief marine phase can be
proposed by looking at Holocene events in the Antarctic Penin-
sula. New evidence in marine cores from the Prince Gustaf Chan-
nel, until recently occupied by part of the Larsen Ice Shelf, indi-
cate that the ice shelf last broke up during the mid- to late-
Holocene (Pudsey and Evans, 2001). If this and other ice shelves
broke up on the Antarctic Peninsula at the same time, there has
been speculation as to whether this could destabilize the inland
glaciers (of the Antarctic Peninsula) and trigger a small sea-level
rise (Cattle et al., 1997). If this speculation is correct, very low-
altitude coastal lakes, emerging (through isostatic recovery) in the
mid- to late-Holocene would be one place where such a change
could be detected. This would be manifested as a brief marine
incursion as a small global sea-level rise temporarily outpaced the
isostatic recovery. If this is the case, such sites offer one potential
analogue for understanding the impacts of modern ice-shelf disin-
tegration on sea level. In order for this lake to be useful in this
respect, a higher-resolution study is required in this lacustrine
zone.
Conclusions
Pup Lagoon is an isolation basin, originated from the sea after
isostatic uplift. Increased productivity and stratified water con-
ditions due to an extensive freshwater input during the retreat of
the Stornes ice sheet are recognized between 5800 and 5500 cal.
yr BP, possibly related to the early to mid-Holocene Hypsither-
mal, detected in ice cores and other palaeoclimatologicalrecords.
Between 2750 and 2200 cal. yr BP conditions are again favour-
able for open-water taxa and high biological productivity, which
may coincide with the late-Holocene warm period detected in
Continental and Maritime Antarctica. A well-defined zone with
brackish-water diatoms and stomatocysts marks the transition
between the marine and the lacustrine phases of the lakes' history.
Chrysophytes are probably among the first colonizers of the newly
exposed coastal lakes, where development of microbial mats is
prevented due to mechanical and/or chemical stress. The isolation
of the lake precedes the emergence of lakes in the Vestfold Hills
with comparable sill heights. After isolation, salinity is relatively
high and fluctuates, followed by a meltwater input resulting in
stable and lower inferred meltwater input. A brief period of
marine incursion following the salinity fluctuations may be related
to local events such as iceberg calving, increased storm activities
or minor sea-level change with sea-level rise temporarily out-
pacing the isostatic recovery.
Acknowledgements
This research is funded by the BAS Signals in Antarctica of Past
Global Changes Program (SAGES), the Federal Office for Scien-
tific, Technical and Cultural Affairs - Belgium Project, Late Quat-
emary Climate History of Antarctic Coastal Environments, a
multiproxy approach (LAQUAN) and the European Commission
through a Framework IV Biotechnology project (MICROMAT
B104-CT98-0040). Elie Verleyen is funded by the Institute for
the Promotion of Innovation by Science and Technology in Fland-
ers. Koen Sabbe is a Senior Research Assistant of the Fund for
Scientific Research (Belgium). Logistical support was provided
by the Australian Antarctic Division (ASAC project 2112). Radio-
carbon dating was supported by the Natural Environment
Research Council Scientific Services and Facilities (734/0498 and
800.0599). Peter Leavitt (University of Regina), Brendan Keely
256 The Holocene 14 (2004)
and Angela Squier (University of York) and NERC Scientific
Services Organic Geochemistry Unit are acknowledged for their
support with pigment identification, the latter two employing
online HPLC-MS. Laboratory support was provided by Pedro
Montiel (BAS), Debbie Hemming and Bruce Terry and analytical
support by Krystyna Saunders. Philippa Noon is thanked for
assistance in the field. Maps were provided by courtesy of the
Australian Antarctic Data Centre and modified by Nick McWil-
liam (BAS). Some background information was extracted from
the 'Visitor's Guide to the Larsemann Hills' which can be found
on the Australian Antarctic Division website. We also thank an
anonymous reviewer and Ben Horton for critically reviewing and
improving the manuscript.
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