Antarctic Science 22(6), 691702 (2010) & Antarctic Science Ltd 2010

doi:10.1017/S0954102010000568

Lichen and moss communities of Botany Bay, Granite Harbour,

Ross Sea, Antarctica
RK2, T.G. ALLAN GREEN3, GERALD MOSER1, STEFAN PANNEWITZ4,
RODNEY D. SEPPELT1, ROMAN TU
LEO G. SANCHO5 and BURKHARD SCHROETER4
1
Australian Antarctic Division, Channel Highway, Kingston, TAS 7050, Australia
Universitat Salzburg, Fachbereich Organismische Biologie, Hellbrunnerstrasse 34, 5020 Salzburg, Austria
3
Biological Sciences, University of Waikato, Private Bag 3105, Hamilton, New Zealand
4
Botanisches Institut, Universitat Kiel, D-24098 Kiel, Germany, and Leibniz Institute for Science and Mathematics Education,
University of Kiel, Olshausenstrasse 62, D-24098 Kiel, Germany
5
Dpto. Biologa Vegetal II, Facultad de Farmacia, Universidad Complutense, 28040 Madrid, Spain
rod.seppelt@aad.gov.au
2

Abstract: Botany Bay is one of the richest sites for lichen and bryophyte biodiversity in continental
Antarctica. A total of 29 lichen, nine moss and one liverwort species have been identified. The most
extensive vegetation occurs on a sheltered raised beach terrace. Vegetation associations are described and
compared to other continental Antarctic localities that also possess a rich vegetation cover. Ordination
analysis clearly indicates the importance of the type of water supply, its regularity, the substrate type, and
particularly in Botany Bay, the influence of nutrients derived from the local bird population in governing
plant distribution and associations. A vegetation map has been produced and can be used as a baseline to
assess vegetation changes over time.
Received 5 April 2010, accepted 9 May 2010

Key words: biodiversity, climate change, cryptogams, vegetation associations, Victoria Land

1994, Seppelt et al. 1988, 1995, Ling & Seppelt 1998, Seppelt
2002) and more rarely from surveys covering larger areas
(Castello & Nimis 1995, Cannone & Seppelt 2008). The
vegetation is composed entirely of cryptogams. Algae and
cyanobacteria dominate the ephemerally wet basins and
stream areas (Howard-Williams & Vincent 1989, HowardWilliams et al. 1989, Vincent et al. 1993, Tang et al. 1997,
Vincent 2000). In localities where at least ephemeral
moisture is plentiful, bryophytes (mosses and one liverwort
Cephaloziella varians) may form extensive patches but
generally their occurrence is patchy, while in dryer areas
their biomass is often exceedingly low. Lichens are found
in a wide variety of habitats, on rock, sand, mosses, weathered
feathers and bones, but rarely is cover or biomass very great.
They may also occur within rocks as endoliths and, in the Dry
Valleys, this is the most common vegetation type over many
square kilometres of the valley floors, predominantly on the
northern faces of suitable moraine boulder debris and country
rock outcrops. Botanical hot spots having substantial
vegetation cover are known from the Canada Glacier,
Taylor Valley and from Botany Bay, Granite Harbour in
southern Victoria Land; Edmonson Point in central Victoria
Land; Beaufort Island in the Ross Sea; Cape Hallett and
Birthday Ridge, northern Victoria Land; and the Windmill
Islands, Wilkes Land. The vegetation of continental Antarctic
regions is a reflection of the rigours of the cold desert
environment where plant distribution is limited primarily by
the availability of free water (Kennedy 1993), although in

Introduction
A detailed knowledge of the composition of the flora and of
the principal vegetation communities in continental Antarctica
(treated here as the main mass of the continent and excluding
the Antarctic Peninsula) is urgently needed as it is realized
that the terrestrial ecosystem may provide one of the earliest
indicators of global climate change, particularly through the
influence of increased temperatures (Petersen & HowardWilliams 2001). Temperatures in some parts of Antarctica are
rising, particularly in the maritime Antarctic (Oppenheimer
1998). Around the northern Antarctic Peninsula, temperature
rises have led to significant losses in the floating ice shelves
and also in significant range extensions southward of the
two indigenous flowering plants (Fowbert & Smith 1994,
Smith 1994) whilst in other areas temperatures are thought to
be falling slowly (Turner 2004, Turner et al. 2005), although
Steig et al. (2009) present evidence for a warming trend that
is much more widespread than previously acknowledged.
Laboratory studies on the soil fauna from the McMurdo Dry
Valleys, Ross Sea region, suggest that drastic changes in
composition or dominance may occur (Doran et al. 2002).
Our knowledge of the vegetation of continental Antarctica
is derived predominantly from a small number of studies in
the vicinity of major stations in coastal regions (e.g. Filson
1966, 1974, Horikawa & Ando 1967, Kuc 196869, Longton
1972, 1973, Seppelt & Ashton 1978, Kappen 1985, Broady
1986, Smith 1988, 1999, Schwarz et al. 1992, Melick et al.
691

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RODNEY D. SEPPELT et al.

some localities such as the Windmill Islands, soil nutrients

derived from relict penguin colonies are also important in
determining abundance (Beyer et al. 2000a, 2000b).
Despite some 60 years of scientific exploration, the
vegetation of the McMurdo region, southern Victoria Land
(77808S, 1601688E), remains relatively poorly understood
or documented. The first attempt at a vegetation classification
was that of Longton (1973) using a modified version of the
classification developed by Gimingham (1967) and Gimingham
& Smith (1970) for the maritime Antarctic. Longton recognized
eleven plant sociations. The overwhelming impression from this
study is one of an impoverished continental Antarctic terrestrial
flora. Only three mosses and eight lichen species were named
and high cover and biomass were found only in dense moss
patches associated with regular meltwater areas. Schwarz et al.
(1992) carried out a detailed study of one of the larger
bryophyte patches, located at the Canada Glacier flush, Taylor
Valley (then Site of Special Scientific Interest SSSI 12, now
Antarctic Specially Protected Area ASPA 131). Although
biomass was locally high with cover up to 100%, only
three mosses Bryum argenteum, B. pseudotriquetrum and
Hennediella heimii were found, the last with aborted
sporophytes (Seppelt et al. 1992). Two lichen species were
subsequently reported growing on the sides of small stones
in the melt stream arising from the side of the glacier and
draining through the moss flush (Green et al. 1992). Species
lists are available for two other sites in the McMurdo Dry
Valleys region, namely Kar Plateau (76856'S, 162820'E)
(Seppelt et al. 1995, 1996) and Beaufort Island (76857'S,
166855'E) (Seppelt et al. 1999). The north end of Beaufort
Island represents an interesting site with a moss bed c. 1 ha
in area with nearly 100% cover composed almost entirely of
one species, Bryum argenteum, and only a very minor cover
of Hennediella heimii. At Cape Hallett (72819'S, 170816'E),
northern Victoria Land, a similar situation occurs with
extensive vegetation dominated by Bryum argenteum and
filamentous green algae occurring on an ephemerally wet
beach terrace, where high nutrient input and atmospheric
ammonium from penguins and skua gulls may limit diversity.
Seppelt & Green (1998) published a bryophyte flora for
southern Victoria Land which included seven moss species
and the liverwort Cephaloziella varians. All these species
can be found at one of the most botanically diverse areas
of continental Antarctica - Botany Bay, Granite Harbour
(778S, 1628E). The first collections of plants from this
region were made by the National Antarctic Discovery
Expedition 190104 (Cardot 1907). The western geological
party of the British Antarctic Expedition of 191013 was
based at Cape Geology, at the northern end of Botany Bay,
but no plant collections appear to have been made
(Schroeter et al. 1993).
We have undertaken extensive fieldwork in the Granite
Harbour region over a number of summer seasons, commencing
with a preliminary survey in December 1989. Detailed studies
on the flora, vegetation, ecology, photosynthetic physiology

of mosses and lichens, and floristic and landscape mapping

began in 1992 and concluded in January 2008. The Botany
BayCape Geology area is now very justifiably designated
as Antarctic Specially Protected Area 154, to conserve and
protect the unique biological richness of the site.
Unless otherwise stated, nomenclature for mosses
follows Ochyra et al. (2008). Nomenclature for lichens
follows vstedal & Lewis Smith (2001). Nomenclature for
liverworts follows Bednarek-Ochyra et al. (2000).

Study area and methods

Site description
Botany Bay and Cape Geology are situated in the southwestern corner of Granite Harbour, southern Victoria Land,
at 162832'5200 E, 77800'1400 S, c. 100 km north-west of Ross
Island. The area consists of raised boulder beach terraces,
weathered rocky steppes and irregular rock platforms
around Cape Geology, rising rapidly to the south to include
a well-defined elevated cirque containing a small ice field.
The bedrock geology at Cape Geology has been described
as a porphyritic grey biotite-granite, with phenocrysts of
orthoclase of reddish colour, casting the weathered rock
with a reddish tinge (Antarctic Treaty Secretariat: http://
www.ats.aq/e/ep_protected.htm). The area faces north and is
well protected from strong winds. The impact of the sun is
also magnified by the sea ice sheet that normally remains in
Granite Harbour until about the end of January so that light
levels are very high. Sensors in the same plane as the lichens
recorded incident PFD (photon flux densities) up to
2800 mmol m-2 s-1, about 40% above normal full sunlight.
Cape Geology and Botany Bay, in particular, are much
warmer than expected and air temperatures can reach
almost 108C with some days in early January being almost
frost free (Schroeter et al. 2010).
The area is extremely rich botanically for such a highlatitude location - it is also one of the richest sites in the whole
of continental Antarctica. There is a high diversity and
abundance of lichens (more than 30 species) and mosses
(nine species), and the structure and development of these
communities are similar to those found 108 of latitude further
north. Some lichen thalli (e.g. Umbilicaria aprina) measure up
to 15 cm diameter. The boulder beach has rich populations of
both epilithic and endolithic lichens. The area is the type
locality for the lichens Buellia frigida (Darbishire 1910) and
Caloplaca coeruleofrigida (Schting & Seppelt 2003). The
area contains by far the most southerly record of an hepatic,
Cephaloziella varians, and the mosses Bryoerythrophyllum
recurvirostrum and possibly Ceratodon purpureus (Seppelt &
Green 1998). There are abundant growths of algae (at least 85
taxa), although the algal flora is not considered particularly
unusual for the locality (Broady 2005).
There are large populations of invertebrates (collembola,
mites, nematodes, rotifers) and the area is the type locality

VEGETATION OF BOTANY BAY

693

Table I. Bryophytes and lichens of the Botany BayCape Geology

region, Granite Harbour, Victoria Land, Antarctica.
HEPATICAE
1
Cephaloziella varians (Gottsche) Steph.
MUSCI
Bryoerythrophyllum recurvirostrum (Hedw.) P.C.Chen
2
Bryum argenteum var. muticum Brid.
Bryum pseudotriquetrum (Hedw.) P.Gaertn., B.Mey. & Scherb.
Ceratodon purpureus (Hedw.) Brid.
3
Didymodon brachyphyllus (Sull.) R.H.Zander
Grimmia plagiopodia Hedw.
Hennediella heimii (Hedw.) R.H.Zander
Schistidium antarctici (Cardot) L.I.Savicz & Smirnova
4
Syntrichia sarconeurum Ochyra & R.H.Zander

Fig. 1. Map of inner Granite Harbour showing the location of

Botany Bay (162832'52''E, 77800'14''S), and other major
named features. Note how Botany Bay is north facing and
well protected from winds coming from the ice shelf
(to the left) and the polar plateau via the McKay Glacier.

for the collembolan Gomphiocephalus hodgsoni. There is a

colony of between 4050 breeding pairs (and numerous nonbreeders) of the south polar skua (Catharacta maccormicki).
These numbers are approximately the same as those present
in 191112 (Taylor 1913, 1916). No other bird species are
known to breed in the Cape Geology area, although Adelie
penguins occasionally come ashore to moult. Nutrient input
from skuas, from meltwater percolating from the ice covered
slopes to the west, and occasionally from the sea when the
sea ice breaks out, plays a significant part in determining the
abundance of the biota, together with water availability from
snowmelt and ephemeral melt streams.
Ecological survey
Relevees were located within the altitudinal range 540 m
above sea level at three primary locations: Botany Bay, Cape
Geology, and the beach terrace near the designated campsite
c. 1 km south-west of Cape Geology (Fig. 1). 186 relevees
were recorded using either 50 cm x 50 cm or 20 cm x 10 cm
quadrats. Species presence was noted and in the larger
quadrats, cover estimates were made visually. For the smaller
quadrats, cover estimates were obtained using point intercepts
at 1 cm intervals with the frame gridded using fine fishing line
and the point intercept being that point at which the lines
crossed in grid corners. Field estimations of cover values
using visual assessment or point quadrats maintained an
accuracy of 1%. Within relevees lichen substrates were
recorded as saxicolous, terricolous or muscicolous.
Botany Bay was also mapped in detail using a building
level to obtain altitude above sea level and plant distribution
mapped using subjective categories which reflected the major
plant type or species present.

LICHENS
Acarospora gwynnii C.W.Dodge & E.D.Rudolph
Amandinea petermannii (Hue) Matzer, H.Mayrhof., & Scheid.
Buellia frigida Darb.
5
Buellia cf. papillata (Sommerf.) Tuck.
6
Buellia subfrigida Mas. Inoue
Caloplaca athallina Darb.
Caloplaca citrina (Hoffm.) Th.Fr.
Caloplaca coeruleofrigida Schting & Seppelt
Caloplaca cf. schofieldii C.W.Dodge
Caloplaca saxicola (Hoffm.) Nordin
Candelariella flava (C.W.Dodge & Baker) Castello & Nimis
7
Carbonea vorticosa (Flrke) Hertel
Lecanora expectans Darb.
Lecanora mons-nivis Darb.
Lecidea andersonii Filson
Lecidea cancriformis C.W.Dodge & G.E.Baker
Lecidella siplei (C.W.Dodge & G.E.Baker) Mas. Inoue
8
Leproloma cacuminum (A.Massal.) J.R.Laundon
Physcia caesia (Hoffm.) Furnr.
Physcia dubia (Hoffm.) Lettau
Rhizocarpon geminatum Korb.
Rhizocarpon geographicum (L.) DC.
Rhizoplaca melanophthalma (Ram.) Leuck. & Poelt
Rhizoplaca cf. priestleyi C.W.Dodge
Sarcogyne privigna (Ach.) A.Massal.
Turgidosculum complicatulum (Nyl.) J.Kohlm. & E.Kohlm.
Umbilicaria aprina Nyl.
9
Xanthomendoza borealis (R.Sant. & Poelt) Schting, Karnefelt &
S.Kondratyuk
Xanthoria elegans (Link.) Th.Fr.
Notes:
1. Cephaloziella varians has previously been referred to as C. exiliflora
(fide Bednarek-Ochyra et al. 2000).
2. Bryum argenteum var. muticum has previously been referred to as
Bryum subrotundifolium (fide Ochyra et al. 2008).
3. Didymodon brachyphyllus has previously been referred to as Didymodon
gelidus (fide Ochyra et al. 2008).
4. Syntrichia sarconeurum has previously been referred to as Sarconeurum
glaciale (fide Ochyra et al. 2008).
5. Buellia cf. papillata, growing on moss, has previously been referred to as
Buellia grimmiae.
6. Buellia subfrigida has previously been referred to as Aspicilia glacialis
(Seppelt et al. 1995) and Hymenelia glacialis (vstedal & Lewis Smith
2001).
7. Carbonea vorticosa has previously been referred to as Lecidea blackburnii
(Seppelt et al. 1995).
8. Leproloma cacuminum has previously been referred to as Lepraria sp.
9. Xanthomendoxa borealis has previously been referred to as Xanthoria
mawsonii (see Lindblom & Schting (2008).

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RODNEY D. SEPPELT et al.

Table II. Buellia frigida sociation composition.

Total average cover

Buellia frigida
Umbilicaria aprina
Xanthoria elegans
Candelariella flava
Xanthomendoza borealis
Rhizoplaca melanophthalma
Physcia caesia
Caloplaca cf. biatorina
Rhizocarpon geminatum
Caloplaca coeruleofrigida
Buellia cf. subfrigida
Rhizocarpon geographicum
Lecidea cancriformis

% Frequency

% Cover

100
77
66
64
50
38.6
26
11
8.6
2.3
2.3
2.3
1.2

54.2
44.9
3.8
4.1
1.32
0.65
0.25
0.43
0.38
0.19
0.08
0.04
0.02
0.01

Data analysis
The classification of data from visual estimates and the 186
relevees by two-way indicator species analysis was
carried out using TWINSPAN 1.0 (Hill 1979) at default
settings. This is a dual technique, providing classification of
samples and species. Groups of samples are characterized
by a characteristic species combination, or at least a group
of differential species (kland 1990). Further ordination
analysis was carried out using a detrended correspondence
analysis (DCA) (Hill & Gauch 1980) using the program
CANOCO 4.0 (ter Braak & Smilauer 1998).

Results
The terrestrial lichen and moss flora of Botany BayCape
Geology comprises one liverwort, nine mosses, and at least
30 lichens (Table I). Several lichen species attributable
to Buellia and Lecidella remain as yet undetermined.
Cyanobacteria, particularly Nostoc, and green algae,
particularly Prasiola crispa, are also abundant in the region.
The classification of the vegetation of the Cape Geology
Botany Bay region includes five lichen-dominated sociations

Table IV. Umbilicaria aprinaRhizocarpon geographicum sociation

composition.

Total average cover

Umbilicaria aprina
Rhizocarpon geographicum
Indeterminate grey crust
Rhizocarpon geminatum
Physcia caesia
Buellia frigida
Candelariella flava

% Frequency

% Cover

83
67
67
33
50
50
16

75.95
20
35
61
3
1
1
2

and two moss dominated sociations within the Crustaceous

Lichen and FolioseFruticose Lichen subformations and
Short Moss Turf and Cushion subformation, with one algal
dominated sociation within the Alga subformation (sensu
Longton 1979). Because of the co-association of both crustose
and foliose lichens over much of the region we have chosen
not to adopt the subformation categories of Longton (1979).
Consequently, the vegetation is classified at the sociation
level.
Antarctic nonvascular cryptogam tundra formation
1. Buellia frigida sociation
This sociation occurs primarily on granitic boulders on the
raised beach terraces, and sheltered sides of massive
country rock risers, steppes and terraces. Water availability
is primarily derived from snow deposits (the boulders can be
almost covered by winter snowfall), by snowmelt from
adjacent snow patches or, very occasionally, by summer
snowfall. In Botany Bay, the south- and south-west-facing
(i.e. towards the cliffs inland) sides of boulders are covered
(almost 100%) by crustose lichens while the north- and northeast-facing (i.e. towards the frozen sea) sides are mostly
devoid of lichen cover. This effect is so strong that when
viewed from the sea ice the boulders in the bay appear to have
no lichen cover at all. Nutrients are derived from snowmelt
and from direct deposition of guano derived from the local
breeding skua gull population. Plant composition in the
70 relevees is summarized in Table II.

Table III. Physcia caesiaBuellia subfrigida sociation composition.

Total average cover

Physcia caesia
Buellia cf. subfrigida
Rhizocarpon geminatum
Umbilicaria aprina
Buellia frigida
Rhizoplaca melanophthalma
Candelariella flava
Rhizocarpon geographicum
Xanthomendoza borealis
Xanthoria elegans
Sarcogyne privigna
Unidentified algae

% Frequency

% Cover

92
95
100
89
79
66
42
36
29
13
5
74

60.5
14.7
7.9
10.8
7.6
6.9
0.54
0.28
0.34
0.17
0.20
0.05
10.9

2. Physcia caesiaBuellia subfrigida sociation

This sociation is predominant in areas of meltwater flow
along ephemeral streams or melt channels and where there is
Table V. Xanthomendoza borealis sociation composition.

Total average cover

Xanthomendoza borealis
Xanthoria elegans
Prasiola crispa
Bryum argenteum
Bryum pseudotriquetrum
Hennediella heimii

% Frequency

% Cover

100
30
70
70
30
8

54
38
0.3
0.8
2.07
0.9
0.31

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VEGETATION OF BOTANY BAY

Table VI. Caloplaca citrina sociation composition.

Total average cover

Caloplaca citrina
Lecanora expectans
Xanthomendoza borealis
Physcia caesia
Xanthoria elegans
Candelariella flava
Caloplaca athallina
Physcia dubia
Rinodina sp.
Nostoc sp.

% Frequency

% Cover

97
83
33
21
4.8
4.8
4.8
9.5
7.1
9.5

19.3
8.4
4.8
5.7
0.28
0.05
0.08
0.08
0.08
0.07
0.67

regular melt seepage over rock. The two dominant species

are largely restricted to these habitats in the region. Physcia
caesia and, to a much lesser extent Physcia dubia, often
form pronounced lines along streams and channels in such a
position that the thalli are regularly covered with water at
peak flow each day. Cyanobacteria and chlorophycean algae
are often present in these habitats but were not identified in
this study. Plant composition in the 38 relevees is
summarized in Table III.
3. Umbilicaria aprinaRhizocarpon geographicum sociation
This sociation is found on sheltered, humid and well-watered
vertical rock faces or along melt seepage or drainage over
stepped ledges and ice fractured country rock. Moisture and
shelter appear to be the principal determining ecological
parameters. Plant composition in the six relevees is
summarized in Table IV.
4. Xanthomendoza borealis sociation
This sociation is found primarily on sandy gravels and coarse
sandy soils and on rock faces in dryer habitats where there is
considerable ornithogenic nutrient input. Often the patches of
gravel and sand are found in shallow depressions, on ledges
and on level areas of country rock where snow and meltwater
collect. Plant composition in the 13 relevees is summarized in
Table V.
5. Caloplaca citrina sociation
This sociation is found on moss cushions and turf and is
dominated by nitrophilous lichens. The moss species are
primarily Hennediella heimii, Bryum pseudotriquetrum
and, to a lesser extent, Bryum argenteum. It is found in
Table VII. Ceratodon purpureus sociation composition.

Total average cover

Ceratodon purpureus
Bryum argenteum
Bryum pseudotriquetrum
Nostoc sp.

% Frequency

% Cover

100
73
47
13

24
21
0.4
0.6
0.4

Table VIII. Bryum argenteumBryum pseudotriquetrum sociation

composition.

Total average cover

Bryum argenteum
Bryum pseudotriquetrum
Hennediella heimii
Ceratodon purpureus
Syntrichia sarconeurum
Xanthomendoza borealis
Prasiola crispa
Nostoc sp.

% Frequency

% Cover

96
75
21
4
4
42
16
8

49
29
13
2.3
0.1
0.13
1.8
1.7
0.1

areas where there is considerable nutrient enrichment from

skua gulls. Moisture is primarily derived from melting
snow. Plant composition in the 42 relevees is summarized
in Table VI.
Short Moss Turf and Cushion subformation
6. Ceratodon purpureus sociation
This sociation is found on moist ledges and on sandy or silty
soils amongst moraine debris and ice rounded boulders,
particularly on the raised beach terrace at Botany Bay. In
terms of water flow this is the wettest part of the area. The
dominant moss is Ceratodon, but Bryum pseudotriquetrum
and, to a lesser extent, other mosses such as Schistidium
antarctici and Bryum argenteum may be found. Plant
composition in the 30 relevees is summarized in Table VII.
7. Bryum argenteumBryum pseudotriquetrum sociation
This sociation is strongly associated with melt streams and
pools, particularly on the raised beach terrace in Botany
Bay. The richest bryophyte communities occur where
moisture is derived directly from melting snowdrifts and
along melt seepage channels and ephemeral streams, or
near temporary shallow pools. Reproduction and dispersal
is almost entirely by asexual propagules and is particularly
prevalent for B. argenteum. Plant composition in the
24 relevees is summarized in Table VIII.
Alga subformation
8. Prasiola crispa sociation
This sociation occurs on nutrient rich sandy soil in habitats
that are sheltered and receive moisture from snowdrifts.
Cover, however, may be low. In nine relevees, cover of
Prasiola was only 1.8%, with total average cover only 2%.
Ordination analysis
The ordination produced some clear groupings that
coincide mainly with those obtained by the sociation

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RODNEY D. SEPPELT et al.

Fig. 2. For caption see next page

analysis (Fig. 2). One difference is the separation into two

parts on the ordination of Buellia frigida sociation 1, one
dominated by B. frigida and the other by Rhizoplaca
melanophthalma, Umbilicaria aprina and several epiphytic

cyanobacterial and algal groups. The spread of this group

indicates that there is more than one habitat on these boulders
which have snowmelt from above and high humidity from
water running between them below, as well as a strong effect

VEGETATION OF BOTANY BAY

697

Fig. 2. a. Two-dimensional ordination of the species and sample plots based on detrended correlation analysis (DCA) of the cover
values of the species. The dimension of the axes is SD-units (standard deviation) so that the variance of the distribution of the
species is shown in this biplot. The position of each sample in the biplot is determined by the species occurring in it. The species
names have been abbreviated to the first four letters of the genus and the first three of the species, the number following the name
classifies the type of occurrence: 7 5 species on cobbles and boulders, 8 5 species on mosses, and 9 5 species on soil. b. As for
a but with the sociations produced by the ordination marked with dotted lines. Red lines are lichen dominated and green lines, moss
dominated. In each case the number corresponds with the sociation number in the text. c. As for a & b but with the form in which
water occurs given for each sociation. Snowmelt means that the water is generated in situ by melting whilst the other categories
indicate the water is flowing or ponding and comes from sources outside the Bay. The dotted lines relate to habitat types and
different associations (e.g. 5, 7) may occupy the same or similar habitat.

from bird fertilization leading to the presence of many

nitrophilous lichen species. A defining feature of this sociation
is that liquid water is rarely present.
The remaining sociations reflect the presence of water,
whether flowing or still, as well as the type of substrate.
Sociation 2 is dominated by Physcia caesia which, in this area
and many other sites along the Ross Sea coast, forms a clear
line along runnels with fluctuating water levels. Typically the
lichen is submerged only at high flows. This contrasts with the
ecology of the species in temperate areas where it is in no way
associated with water flows. Sociation 6, which is bryophyte
dominated, falls in the same location on the ordination and is
dominated by Ceratodon purpureus. In this case the most
probable difference is the less stable substrate. Flowing water
over unstable but not steep substrates is the location of the
second bryophyte dominated sociation, sociation 7, with high
cover of Bryum pseudotriquetrum and B. argenteum and this
is a very typical occurrence in the Ross Sea region when there
is the correct combination of an excellent, regular, slow
flowing water supply together with good shelter. Flowing

water over a sheltered but well lit rock surface are the
apparent conditions for sociation 3, dominated by U. aprina
and Rhizocarpon geographicum. Again, for the latter species,
this is not the expected habitat from its alpine ecology.
Sociation 5, characterized by a dominance of Caloplaca
citrina, with the lichens growing primarily on mosses. This is
a habitat that is marginal to the main water flows and one in
which the lichens do not suffer flooding but clearly obtain a
good water supply from the moss beneath them. The presence
of nutrients from birds and somewhat drier conditions seems
to determine the occurrence of this sociation. Sociation 4,
characterized by the presence of Xanthomendoza borealis,
occurs typically on periodically wetted, but often unstable
rock ledges with a strong bird nutrient influence. The alga
Prasiola crispa can occur in similar sites, especially those
with pooled water.
The analysis clearly indicates the importance of the type
of water supply, its regularity, the substrate type, and
particularly in Botany Bay, the influence of nutrients
derived from the local bird population.

698

RODNEY D. SEPPELT et al.

Fig. 3. Vegetation map of Botany Bay, Granite Harbour, southern Victoria Land. Grid interval 5 10 m.

Botany Bay vegetation map

Vegetation zonation and topography in Botany Bay are
shown in Fig. 3. Shape and topography of the seaward
margin is influenced by annual events relating to sea ice.
With breakout of the sea ice, there is opportunity through
ice push to shape the topography. However, while creating
disturbance at the seaward margin of the bay, this has little
influence on the vegetation. Meltwater passing over areas
of moss, particularly B. argenteum, results in the release
and subsequent deposition of deciduous apical shoot
propagules along the melt channels. Establishment of the
moss colonies in the sand and boulders at the shore is
almost impossible as the habitat is too unstable.

Discussion
Since we first visited the area and began the detailed
mapping and ecological surveys there has been little
obvious change in plant distribution patterns. However,
under the influence of global warming trends, increased
water flow through or over the vegetation may lead to

change. Drainage lines may become wetter or drier,

depending on the direction of melt flow over the raised
beach terrace, leading to changes in the distribution pattern
of the vegetation. Our detailed vegetation map can be used
as a baseline reference to monitor such changes.
In the Windmill Islands, the only other continental
Antarctic region where vegetation mapping on a large scale
has been undertaken, there has been appreciable change in
the vitality of moss beds over the last 10 years (R. Seppelt,
S. Robinson and J. Wasley, personal observations) due to
changes in the moisture regime of the habitat. At the Canada
Glacier, Taylor Valley, increased summer meltwater flow
over the large moss flush has resulted in pronounced lateral
spread of the moss. If the warming trends continue to
influence continental Antarctic regions we may expect
significant change to the extent of the moss cover. Careful
and accurate mapping of the vegetation in selected areas such
as these (particularly Canada Glacier, Botany Bay, Beaufort
Island and Cape Hallett in Victoria Land, the Windmill
Islands region, Wilkes Land) has provided an important tool
for assessing the impacts of climatic change on the continental
Antarctic terrestrial ecosystem.

VEGETATION OF BOTANY BAY

The surveys carried out in this investigation fully confirm

that Botany Bay is one of the richer sites for lichen and
bryophyte biodiversity in continental Antarctica. The total of
29 lichen, nine moss and one liverwort species is practically
identical to the numbers reported for Wilkes Land by Smith
(1988), a site that is about 11 degrees latitude further north
(668S compared to 778S for Botany Bay). Around thirty
lichen species are also reported for Casey and Davis
stations, also at about 668S latitude (Green et al. 2007). It
is probable that Edmonson Point, Terra Nova Bay, has
higher numbers of species but the 56 species reported from
there (Castello & Nimis 1995) include lichens from several
sites further north. The similarity in total species numbers
across the whole of continental Antarctica suggests that the
cline in biodiversity seen as one moves south along the
Antarctic Peninsula (Peat et al. 2007) does not continue
over continental regions. Further indicators of the special
nature of Botany Bay are the southernmost records
in Antarctica for a liverwort, Cephaloziella varians, for
the lichen Turgidosculum complicatulum, for the moss
Bryoerythrophyllum recurvirostrum and probably also for
Ceratodon purpureus. Most are about three degrees latitude
further south than the nearest record to the north in the
Terra Nova Bay area.
The results of the vegetation analysis also agree in the
main with those of Smith (1988) for Wilkes Land and
Cannone & Seppelt (2008) and Castello & Nimis (1995) for
central and northern Victoria Land, respectively. The latter
two studies covered a larger geographic spread than this
study and the level of agreement suggests that there is
limited diversity of vegetation types in the Ross Sea region.
Our groups 1 and 2, B. frigida sociation on rocks and the
P. caesia dominated fluvial sociation, agree with 2A and 2B
of Cannone & Seppelt (2008). Similarly, our groups 3 and 4
correspond to group 3, our group 5 to group 4, and the
bryophyte dominated groups 6 and 7 to groups 6.1 and 6.2
of Cannone & Seppelt (2008). The main determinants of
the occurrence of the different sociations appears to be the
source of water, whether solely from snowmelt or snowfall,
or from some form of melt stream. Sites influenced primarily
by snow cover are lichen dominated while those influenced by
meltwater are either mixed bryophyte/lichen or, in the wetter
places, bryophyte dominated. Sociations are also influenced
by the regularity and speed of water flow, and the type of
substrate, especially whether it is loose gravel or solid rock.
Some species show different ecologies in Antarctica
compared to their temperate or alpine habitats or even
from elsewhere in Antarctica. Rhizocarpon geographicum
is typically found in Botany Bay and nearby localities
where water flows slowly over a near vertical solid rock
surface. Physica caesia is found as a distinct line along
watercourses at a level where it is regularly submerged
each day at peak summer melt flow. Neither species
is associated with flowing water elsewhere. The moss
C. purpureus is typically reported from drier sites in

699

flushes, such as at Edmonson Point (Smith 1999), but it

occurs in the wettest habitats at Botany Bay.
Possibly the greatest difference between Botany Bay and
the other sites to the north is the complete absence of Usnea
species. This is not easy to explain as U. antarctica is
reported from Kar Plateau, just across Granite Harbour
from Cape Geology, and U. sphacelata is reported from
848S at Mount Kyffin, in the southern Ross Sea. Abundant
populations of U. sphacelata and U. antarctica also occur
on Ross Island at sites that are visible from Cape Geology.
In contrast, the specimens of Umbilicaria aprina at Botany
Bay are some of the larger specimens reported in Antarctica
and were reported to be as large as dinner plates by Scotts
western geological party in 1912 (Schroeter et al. 1993).
The generalization of Smith (1988) that three main
groups occur, one dominated by lichens, one by bryophytes
and a third in which lichens are epiphytic on bryophytes
seems to hold. There seems little doubt that the extremely
moderate summer climate in Botany Bay, the excellent
water supply and nutrient input from the skua colony are
the main determinants of the rich vegetation. The north
facing aspect, the enhanced radiation from the almost everpresent sea ice and protection from strong winds afforded
by cliffs to the south and west, means that very warm
conditions occur for about two months each year. At the
beginning of January there are a small number of days that
are almost frost free with maximal air temperatures close to
108C. The presence of the corrie to the south and above the
Bay, and of a perched ice field above, ensures a steady
source of meltwater in summer. Winter snowfalls appear
to also be regular but not excessive, only about 40 cm
depth in total. Shallow snowfalls are advantageous in this
area as late lying snow acts as an insulator, keeping the
ground cold and delaying the onset of active metabolism
(Pannewitz et al. 2003) until air temperatures are warmer
and water supply more reliable. Even so, it is dangerous to
assume that a high visible biomass also means high
productivity. The studies by Kappen et al. (1998) clearly
show that B. frigida thalli on boulders are only active for a
small number of hours each year. At least for this species,
growth rates are exceedingly variable, ranging from 1.0 mm
per 100 years in the Taylor Valley to 6.9 mm per 100 years
at Cape Hallett (728S) (Sancho et al. 2007).
We are now able to compare descriptions from three sites
at about 778S latitude: Canada Glacier flush, an extensive
area of bryophyte dominated vegetation in lower Taylor
Valley (Schwarz et al. 1992); Beaufort Island, an area of
c. 1 ha of almost pure B. argenteum on a north facing slope
backed by a glacier wall and with an extensive skua colony
(Seppelt et al. 1999), and Botany Bay (present study). A
comparison on the basis of lichens is not possible as they
are almost absent except at Botany Bay. The bryophytes are
interesting in that there are only three species at Canada
Glacier, ten at Botany Bay (one liverwort) and two (one
dominant moss and one very minor) on Beaufort Island.

700

RODNEY D. SEPPELT et al.

A possible explanation for these differences could be the level

of external nutrition. There is almost no external nutrient
supply at Canada Glacier except that brought by dust or
glacier meltwater. At Botany Bay there is a moderate input
from the small skua colony and at Beaufort Island a
significant input from the extensive skua colony. It appears
that there may be an optimal nutrient level for bryophyte
biodiversity and that too little, or especially an excess, will
lead to lowered species numbers. It is also of note that at Cape
Hallett, where there is an extensive bryophyte flush adjacent
to skua nests and a very large Adelie penguin colony, there is
only one moss species, B. argenteum, in the main flush area.
This is an equivalent situation to Beaufort Island and suggests
that only B. argenteum can take advantage of these high
nutrient zones. In other coastal areas of continental Antarctica,
such as the Syowa Coast (408E) (Kanda 1981, 1987, Kanda &
Inoue 1994), B. argenteum is the dominant moss near to the
shore, but to the east, between 508E and 1608E, this species
appears to be absent, being replaced by B. pseudotriquetrum.
Despite the growing interest in the possible effects of
climate change on Antarctic vegetation it seems that we are a
long way from being able to successfully predict vegetation
shifts in the Ross Sea region. In the maritime Antarctic
Peninsula a clear biodiversity cline exists and must depend on
a climatic variable, probably water availability. However, in
the Ross Sea region there is little evidence of a clear cline,
similar vegetation types seem to exist all along the coast and
even to more distant sites, like Wilkes Land. This is almost
certainly a result of the desert conditions with suitable habitats
only becoming available where a regular and substantial water
supply is available. In the Windmill Islands (668S) there has
been a marked rapid change in the vitality of the mossdominated areas (Melick et al. 1994, Melick & Seppelt 1997,
Wasley et al. 2006, Seppelt et al. unpublished observations)
with an increase in abundance of epiphytic lichen cover
and moribund moss being associated with a drying of the
vegetation. Distributional changes, particularly in algae and
mosses and probably relating to altered water flows have also
been reported from Cape Hallett (Brabyn et al. 2005, 2006.
Some properties of the vegetation, like lichen growth rate
(Sancho et al. 2007), DNA damage in mosses following
greater exposure to higher levels of incident UV radiation
(Turnbull & Robinson 2009), and changes in flavonoid
chemistry resulting from changes in atmospheric ozone levels
(Ryan et al. 2009) do seem to be possible indicators of climate
change. However, without an assessment of changes in water
availability the actual vegetation, both its structure and
biodiversity, does not appear to offer sufficient sensitivity to
be of use as an indicator at the moment.
Acknowledgements
We thank Antarctica New Zealand for provision of logistic
support and the University of Waikato, through their Antarctic
Research Program, for their continuing support for Antarctic

research over many years. BS and SP gratefully acknowledge

financial support from the Deutsche Forschungsgemeinschaft
(SCHR 473/4-3). TGAG was supported by a Ramon y Cajal
Fellowship at Vegetal II, Farmacia, Universidad Complutense,
Madrid, Spain, and by FRST grant: Understanding, valuing
and protecting Antarcticas unique terrestrial ecosystems:
Predicting biocomplexity in Dry Valley ecosystems, during
the writing of this paper.
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