Lichen and moss communities of Botany Bay, Granite Harbour, Ross Sea, Antarctica

Antarctic Science 22(6), 691–702 (2010) & Antarctic Science Ltd 2010 doi:10.1017/S0954102010000568

Lichen and moss communities of Botany Bay, Granite Harbour,Ross Sea, Antarctica

RODNEY D. SEPPELT1, ROMAN TURK2, T.G. ALLAN GREEN3, GERALD MOSER1, STEFAN PANNEWITZ4,LEO G. SANCHO5 and BURKHARD SCHROETER4

1Australian Antarctic Division, Channel Highway, Kingston, TAS 7050, Australia2Universitat Salzburg, Fachbereich Organismische Biologie, Hellbrunnerstrasse 34, 5020 Salzburg, Austria

3Biological Sciences, University of Waikato, Private Bag 3105, Hamilton, New Zealand4Botanisches Institut, Universitat Kiel, D-24098 Kiel, Germany, and Leibniz Institute for Science and Mathematics Education,

University of Kiel, Olshausenstrasse 62, D-24098 Kiel, Germany5Dpto. Biologıa Vegetal II, Facultad de Farmacia, Universidad Complutense, 28040 Madrid, Spain

[email protected]

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

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 & Howard-

Williams 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 1968–69, Longton

1972, 1973, Seppelt & Ashton 1978, Kappen 1985, Broady

1986, Smith 1988, 1999, Schwarz et al. 1992, Melick et al.

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, Howard-

Williams 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

691

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

(77–808S, 160–1688E), 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 1901–04 (Cardot 1907). The western geological

party of the British Antarctic Expedition of 1910–13 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

Bay–Cape 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 Øvstedal & 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 south-

western corner of Granite Harbour, southern Victoria Land,

at 162832'5200E, 77800'1400S, 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 high-

latitude 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 (Søchting & 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

692 RODNEY D. SEPPELT et al.

for the collembolan Gomphiocephalus hodgsoni. There is a

colony of between 40–50 breeding pairs (and numerous non-

breeders) of the south polar skua (Catharacta maccormicki).

These numbers are approximately the same as those present

in 1911–12 (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 5–40 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.

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.

Table I. Bryophytes and lichens of the Botany Bay–Cape Geology

region, Granite Harbour, Victoria Land, Antarctica.

HEPATICAE1Cephaloziella varians (Gottsche) Steph.

MUSCI

Bryoerythrophyllum recurvirostrum (Hedw.) P.C.Chen2Bryum argenteum var. muticum Brid.

Bryum pseudotriquetrum (Hedw.) P.Gaertn., B.Mey. & Scherb.

Ceratodon purpureus (Hedw.) Brid.3Didymodon brachyphyllus (Sull.) R.H.Zander

Grimmia plagiopodia Hedw.

Hennediella heimii (Hedw.) R.H.Zander

Schistidium antarctici (Cardot) L.I.Savicz & Smirnova4Syntrichia sarconeurum Ochyra & R.H.Zander

LICHENS

Acarospora gwynnii C.W.Dodge & E.D.Rudolph

Amandinea petermannii (Hue) Matzer, H.Mayrhof., & Scheid.

Buellia frigida Darb.5Buellia cf. papillata (Sommerf.) Tuck.6Buellia subfrigida Mas. Inoue

Caloplaca athallina Darb.

Caloplaca citrina (Hoffm.) Th.Fr.

Caloplaca coeruleofrigida Søchting & Seppelt

Caloplaca cf. schofieldii C.W.Dodge

Caloplaca saxicola (Hoffm.) Nordin

Candelariella flava (C.W.Dodge & Baker) Castello & Nimis7Carbonea vorticosa (Flørke) 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. Inoue8Leproloma 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.9Xanthomendoza borealis (R.Sant. & Poelt) Søchting, 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 (Øvstedal & 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 & Søchting (2008).

VEGETATION OF BOTANY BAY 693

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 (Økland 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 Bay–Cape

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

and two moss dominated sociations within the Crustaceous

Lichen and Foliose–Fruticose 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 north-

east-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.

2. Physcia caesia–Buellia subfrigida sociation

This sociation is predominant in areas of meltwater flow

along ephemeral streams or melt channels and where there is

Table II. Buellia frigida sociation composition.

% Frequency % Cover

Total average cover 54.2

Buellia frigida 100 44.9

Umbilicaria aprina 77 3.8

Xanthoria elegans 66 4.1

Candelariella flava 64 1.32

Xanthomendoza borealis 50 0.65

Rhizoplaca melanophthalma 38.6 0.25

Physcia caesia 26 0.43

Caloplaca cf. biatorina 11 0.38

Rhizocarpon geminatum 8.6 0.19

Caloplaca coeruleofrigida 2.3 0.08

Buellia cf. subfrigida 2.3 0.04

Rhizocarpon geographicum 2.3 0.02

Lecidea cancriformis 1.2 0.01

Table III. Physcia caesia–Buellia subfrigida sociation composition.

% Frequency % Cover



Buellia cf. subfrigida 95 7.9

Rhizocarpon geminatum 100 10.8

Umbilicaria aprina 89 7.6

Buellia frigida 79 6.9

Rhizoplaca melanophthalma 66 0.54

Candelariella flava 42 0.28

Rhizocarpon geographicum 36 0.34



Sarcogyne privigna 5 0.05

Unidentified algae 74 10.9

Table IV. Umbilicaria aprina–Rhizocarpon geographicum sociation

composition.

% Frequency % Cover


Umbilicaria aprina 83 20

Rhizocarpon geographicum 67 35

Indeterminate grey crust 67 61

Rhizocarpon geminatum 33 3

Physcia caesia 50 1

Buellia frigida 50 1

Candelariella flava 16 2

Table V. Xanthomendoza borealis sociation composition.

% Frequency % Cover

Total average cover 54

Xanthomendoza borealis 100 38


Prasiola crispa 70 0.8

Bryum argenteum 70 2.07

Bryum pseudotriquetrum 30 0.9

Hennediella heimii 8 0.31


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 aprina–Rhizocarpon 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

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 argenteum–Bryum 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

Table VI. Caloplaca citrina sociation composition.

% Frequency % Cover


Caloplaca citrina 97 8.4

Lecanora expectans 83 4.8



Xanthoria elegans 4.8 0.05

Candelariella flava 4.8 0.08

Caloplaca athallina 4.8 0.08

Physcia dubia 9.5 0.08

Rinodina sp. 7.1 0.07

Nostoc sp. 9.5 0.67

Table VII. Ceratodon purpureus sociation composition.

% Frequency % Cover


Ceratodon purpureus 100 21

Bryum argenteum 73 0.4

Bryum pseudotriquetrum 47 0.6

Nostoc sp. 13 0.4

Table VIII. Bryum argenteum–Bryum pseudotriquetrum sociation

composition.

% Frequency % Cover


Bryum argenteum 96 29

Bryum pseudotriquetrum 75 13

Hennediella heimii 21 2.3

Ceratodon purpureus 4 0.1

Syntrichia sarconeurum 4 0.13


Prasiola crispa 16 1.7

Nostoc sp. 8 0.1


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

Fig. 2. For caption see next page


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.

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.


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.

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


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

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 Scott’s

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 ever-

present 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.


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 moss-

dominated 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 Antarctica’s unique terrestrial ecosystems:

Predicting biocomplexity in Dry Valley ecosystems, during

the writing of this paper.

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Lichen and moss communities of Botany Bay, Granite Harbour, Ross Sea, Antarctica

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