Algological Studies 126 173–208 Stuttgart, April 2008

Cyanobacteria in microbial mats of

Antarctic lakes (East Antarctica) –
A microscopical approach

ARNAUD TATON1, 2, LUCIEN HOFFMANN3 and ANNICK WILMOTTE2*

1 Institut de Botanique, Université de Liège, Liège, Belgium

2 Centre d’Ingénierie des Protéines, Université de Liège, Liège, Belgium
3 Department ‘Environment and Agro-biotechnologies’, Centre de Recherche
Public-Gabriel Lippmann, Belvaux, Grand-duchy of Luxembourg

With 13 figures and 2 tables

Abstract: The cyanobacterial diversity in microbial mats of 56 lakes located in

the Larsemann Hills and Bølingen Islands (East Antarctica) was studied by light
microscopy. Thirty taxa belonging to 13 genera of Chroococcales, Oscillatoriales
and Nostocales were described. Photomicrographs of the different taxa are pro-
vided and their taxonomical assignment as well as their ecology and distribution
are discussed. In addition, a semi-quantitative estimation of the abundance of the
different morphotypes in the samples was realized and correlated with the mat
physiognomy.

Key words: Antarctic lakes, biogeography, cyanobacteria, ecology, microbial

mats, morphological diversity, taxonomy

Introduction
Microbial mats are an important component of Antarctic lakes and ponds.
They are usually dominated by cyanobacteria and often constitute the larg-
est accumulation of cyanobacteria in Antarctic biotopes (VINCENT 2000b).
The morphological diversity of cyanobacteria in Antarctic microbial
mats has been studied since 1911 (WEST & WEST 1911). Lakes and ponds
in the region of the McMurdo station were mainly investigated: Ross Island
(WEST & WEST 1911, BROADY & KIBBLEWHITE 1991), McMurdo Ice Shelf

* Corresponding author.

DOI: 10.1127/1864-1318/2008/0126-0173 1864-1318/08/0126-173 $ 9.00

© 2008 E. Schweizerbart’scheVerlagsbuchhandlung, D-70176 Stuttgart

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 174 A. TATON et al.

(NADEAU et al. 2001), McMurdo Sound (FRITSCH 1912, PRESCOTT 1979), the
Ross Sea Region and the Dry Valleys (Southern Victoria Land) (PARKER et
al. 1975 according to PRESCOTT 1979, WHARTON et al., 1983, SIMMONS et al.,
1993) (Fig. 1). Furthermore, mat-forming cyanobacteria were also studied
from lakes and ponds of Inexpressible Island (Northern Victoria Land)
(FUMANTI et al. 1997), from lakes of Signy Islands (South Orkney Islands)
(PRIDDLE & BELCHER 1982) and lakes and pools of Hope Bay (Peninsula)
(VINOCUR & PIZARRO 1995).
To date, few data are available concerning the cyanobacterial diversity
of microbial mats in East Antarctica. The physical, chemical and microbial
community characteristics of several lakes of the Larsemann Hills were
studied by ELLIS-EVANS et al. (1996), who have observed a large variety
of mat types with different structures and compositions depending on the
lake depth, water conductivity and the zonation within the lake. More
recently, the structure and composition of mats in 56 lakes and ponds of
the Larsemann Hills (SABBE et al. 2004) and their pigment composition
(HODGSON et al. 2004) were investigated. The mat physiognomy and the
pigment composition were strongly correlated to lake depth (SABBE et
al. 2004, HODGSON et al. 2004). However, up to date, no morphological
description or list of the cyanobacterial taxa was provided.
In the present study, the cyanobacterial diversity of 56 lakes and ponds
of the Larsemann Hills investigated by the latter authors was analyzed
in order to improve our knowledge about the cyanobacterial diversity
of these mats. Morphological descriptions and photomicrographs of the
different cyanobacterial morphotypes are provided and their taxonomical
assignment as well as their ecology and distribution are discussed.

Materials and methods

Study area

The Larsemann Hills (69°23’S, 76°53’E) located in the Prydz Bay region,
constitute a major ice-free area in continental East-Antarctica of around
50 km2 (HODGSON et al. 2001, VERLEYEN et al. 2004). The Larsemann
Hills region consists of two main peninsulas, the western Stornes and the
eastern Broknes/Mirror, together with a number of scattered offshore is-
lands. The Bølingen Islands form a smaller largely ice-free archipelago,
twenty-five kilometers to the west south west of the Larsemann Hills.
More than 150 freshwater lakes are found in the hills (GILLIESON et al.
1990) ranging from small ephemeral ponds to large water bodies. Some of
these water bodies are briefly ice-free or partially ice-free in the summer
months when their water temperature rapidly increases. For the remain-
ing 8–10 months of the year, they are covered with ca. 2 m of ice. Lake
systems are connected to the coast by large, steep-sided V-shaped valleys

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 Cyanobacteria in mats of East Antarctic lakes 175

Fig. 1. Locations of the different regions mentioned in the text: EL, Enderby
Land; MAR, Maritime Antarctica; MBL, Marie Byrd Land; MRL, Mac Robertson
Land; NVL, Northern Victoria Land; PEL, Princess Elizabeth Land; PEN, Penin-
sula; QML, Queen Maud Land; RSR, Ross Sea Region; SVL, Southern Victoria
Land; WL,Wilkes Land.

that dissect the area as well as by streams that flow continuously during
the summer months.

Biological sampling

In the Antarctic summer of 1997–1998, 51 lakes were sampled in the Larse-

mann Hills and five lakes in the Bølingen Islands. Lakes were chosen in or-
der to represent the whole range in lake morphometry, hydrological char-
acteristics and salinity in the region (GILLIESON et al. 1990, ELLIS-EVANS et
al. 1998).
Samples of benthic microbial mats were collected from the deepest parts
of the lakes using a Glew surface sediment corer enabling to accurately

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 176 A. TATON et al.

Table 1. Studied lakes

Lake number Lake name Longitude – Latitude Cyanobacteria
L72 Lake Nella 76°22’E – 69°24’S +
L35 Crater Lake 76°11’E – 69°23’S
L34 Kirisjes Pond 76°09’E – 69°22’S +
L12 Long Lake 76°07’E – 69°24’S
LBU Lake Burgess 76°07’E – 69°25’S +
L36 – 76°13’E – 69°25’S
BBA – 76°05’E – 69°27’S +
L57 Progress Lake 76°24’E – 69°24’S
L51 Lake Cameron 76°21’E – 69°24’S
L71 Sarah Tarn 76°23’E – 69°23’S +
L59 Moore Lake 76°21’E – 69°24’S +
L67 – 76°21’E – 69°23’S +
L74 Discussion Lake 76°22’E – 69°23’S +
L63 – 76°18’E – 69°23’S +
L60 – 76°20’E – 69°23’S +
L8 – 76°05’E – 69°09’S +
L7 – 76°05’E – 69°09’S
L10 Lake Heidi 76°06’E – 69°24’S +
L13 – 76°07’E – 69°24’S +
L14 – 76°07’E – 69°24’S +
L23 Pup Lagoon 76°03E – 69°25’S +
LGR – 76°11’E – 69°24’S +
L49 – 76°16’E – 69°24’S +
L44 – 76°17’E – 69°24’S +
L43 – 76°15’E – 69°23’S +
L69 No Worries Lakes 76°23’E – 69°22’S
L70 Lake Reid 76°23’E – 69°23’S +
L68 Heart Lake 76°23’E – 69°23’S +
BAL Lake Alanna 75°55’E – 69°28’S
LJA Lake Jack 76°06’E – 69°25’S +
L18 Lake Spate 76°07’E – 69°25’S +
L1 Lake Anna 76°17’E – 69°23’S +
L73 – 76°23’E – 69°24’S +
LSP – 76°02’E – 69°25’S +
L42 – 76°15’E – 69°23’S +
BST – 75°50’E – 69°27’S
LPR – 76°23’E – 69°23’S
BFI Firelight Lake 75°45’E – 69°31’S
L53b – 76°23’E – 69°24’S +
LPS – 76°23’E – 69°23’S +
L52 Lake Bruehwiler 76°21’E – 69°24’S +
L52b – 76°21’E – 69°24’S +
L66 – 76°20’E – 69°24’S +
L65 – 76°19’E – 69°24’S +
LG2 – 76°19’E – 69°23’S +
L64 – 76°18’E – 69°23’S +
LSN – 76°18’E – 69°23’S +

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 Cyanobacteria in mats of East Antarctic lakes 177

Lake number Lake name Longitude – Latitude Cyanobacteria

L61 – 76°19’E – 69°22’S +
BSU Sunset Lake 75°45’E – 69°31’S +
L58 Lake Sibthorpe 76°21’E – 69°24’S
LFO – 76°20’E – 69°21’S +
LMA – 76°19’E – 69°21’S +
L59b – 76°21’E – 69°24’S +
LG1 – 76°19’E – 69°23’S +
LG4 – 76°19’E – 69°23’S
L64b – 76°18’E – 69°23’S +

sample the top 0.5 to 1 cm of the mats. The sampled layers thus included
several years of mat growth and therefore accounted (as far as possible)
for inter-annual variations in winter ice duration, thickness, transparency
and snow cover. When no ice cover was present (lakes < 2 m deep), samples
were taken between 0.5–1 m in the littoral zone. All samples were frozen
and stored in the dark until they were transferred in a 4% formaldehyde
solution. The abiotic characteristics of the sampled lakes were published by
SABBE et al. (2004). The lake numbers, names, latitudes and longitudes are
listed in table 1.

Analysis of materials

The cyanobacterial taxa were observed with a Wild MS-20 microscope

equipped with a screw micrometer and a camera Lucida or with a Zeiss
Universal microscope equipped with a digital camera. The diacritical mor-
phological traits used for morphological species descriptions were the cell
shape for both intercalary and end cells; the width and length of intercalary
cells; the presence or absence of constriction at the cross wall, of necridic
cells, and of a sheath, color of the sheath, the number of trichomes per fila-
ment, the presence or absence of heterocysts, the width and length of het-
erocysts. For each biometrical character, at least 30 measurements were ob-
tained from cells, heterocysts, and filaments sampled at random. Measure-
ments were realized with a screw micrometer or using the software ImageJ
[http://rsb.info.nih.gov/ij/] from pictures. The small unicellular types as well
as the thin Oscillatoriales were particularly difficult to identify directly from
the field samples. Therefore, for each sample, pictures of taxa appearing to
be different in light microscopy were taken. Based on these pictures, cell
measurements were realized and morphological diacritical traits were used
to distinguish the different morphotypes. The taxonomic works of GEITLER
(1932), ANAGNOSTIDIS & KOMÁREK (1985), and KOMÁREK & ANAGNOSTIDIS
(1989, 1998, 2005), as well as descriptions of Antarctic cyanobacteria were
used. In accordance with KOMÁREK’s (1999) species concept (discussed be-
low) for Antarctic taxa we have used “cf.” for the species found for the first

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 178 A. TATON et al.

Table 2: Morphotypes and corresponding taxonomic assignments

Morphotype Taxonomic assignment
U05–U06 Aphanocapsa cf. holsatica (LEMMERMANN) CRONBERG et KOMÁREK
1994
U09 Aphanocapsa cf. hyalina (LYNGBYE) HANSGIRG 1892
U03 Aphanothece cf. elabens (BRÉBISSON in MENEGHINI) ELENKIN 1938
U04 Aphanothece cf. microscopica NÄGELI 1849
U01 Aphanothece sp.
ASru Asterocapsa sp. (Gloeocapsa cf. rupestris KÜTZING 1845)
U08 Cf. Chamaesiphon
CHsu Chamaesiphon cf. subglobosus (ROSTAFINSKI) LEMMERMANN 1910
U07 Chlorogloea sp.
CHde Chondrocystis cf. dermochroa (NÄGELI) KOMÁREK et ANAGNOS-
1995
TIDIS

GLal Gloeocapsa cf. alpina (NÄGELI) BRAND 1900

GLco Gloeocapsa cf. compacta KÜTZING 1845
GLsa Gloeocapsa cf. sanguinea (AGARDH) KÜTZING 1843
U02 Rhabdonema sp.
LPP04 Cf. Jaaginema sp.
LPP02 Leptolynbya sp.
OSs1 Phormidium autumnale (AGARDH) TREVISAN ex GOMONT 1892
LYs1 Phormidium murrayi (W. et G. S. W EST ) A NAGNOSTIDIS et
KOMÁREK 1988
LPP01–LPP06 Phormidium priestleyi FRITSCH 1917
LPP08 Pseudanabaena cf. amphigranulata (VAN GOOR) ANAGNOSTIDIS
2001
LPP03 Pseudanabaena cf. tremula CASAMATTA et JOHANSEN 2005
LPP05-LPP07 Pseudanabaena frigida (FRITSCH) ANAGNOSTIDIS 2001
SCs1 Schizothrix cf. lacustris A. BRAUN ex GOMONT 1892
SCs3 Schizothrix cf. simplicior SKUJA 1964
SCs2 Schizothrix cf. tenuis VORONICHIN 1923
CAs1 Calothrix cf. parietina (NÄGELI) THURET ex BORN. et FLAH. 1886
COsc Coleodesmium cf. scottianum WELSH 1965
DIsp Dichothrix sp.
NOsp Nostoc sp.
PEin Petalonema cf. involvens (A. BRAUN) ex MIGULA

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 Cyanobacteria in mats of East Antarctic lakes 179

time in Antarctica and for those for which the ecological range given in the
literature does not correspond to the ecology of the type locality.
A semi-quantitative estimation of the relative abundance of each taxon
in the studied samples was realized following a method derived from
GOLUBIC (1967). The surface occupied by each taxon was determined by
the superposition of a grid on the microscopic field. The “countings” were
made using two magnifications (500 x and 1260 x). The grid meshes and the
microscopic fields were 2 µm and 1258 µm2 at 500 x and 5 µm and 8000 µm2
at 1260 x, respectively. For each sample, three replicate preparations and
15 fields/preparation at both magnifications were examined. The average
surfaces occupied by each taxon were then calculated on the basis of the 90
examined fields.

Results and discussion

Descriptions

Fifty-six samples were observed in light microscopy, of which 20 mostly

contained dead cells, empty sheaths or appeared mostly mineral. Finally, 43
samples contained enough cyanobacteria to study their community compo-
sition. Thirty taxa corresponding to 33 morphotypes belonging to 13 genera
of Chroococcales, Oscillatoriales and Nostocales were distinguished. For
some taxa, two morphotypes were found. For each morphometrical feature,
the average and standard deviation as well as the minimum and maximum
dimensions are given. The lake numbers were used in the occurrence field
(see table 1 for the lake names and locations). Morphotypes and corre-
sponding taxonomical assignments are listed in table 2.

Chroococcales

The small Chroococcales were very difficult to identify in the field samples.
Therefore, for each sample, pictures of each cyanobacterium appearing to
be different in light microscopy were taken. Two main groups were dis-
tinguished, the first one possessed spherical cells and the second one cells
longer than wide. Based on the pictures, cell measurements were realized
(Figs 2 to 4) and morphological diacritical traits determined to distinguish
the different morphotypes.

Aphanocapsa cf. holsatica (LEMMERMANN) CRONBERG ET KOMÁREK 1994

B a s i o n y m : Clathrocystis holsatica LEMMERMANN 1903

S y n o n y m : Microcystis holsatica (LEMMERMANN) LEMMERMANN 1907
M o r p h o t y p e : U05

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 180 A. TATON et al.

Fig. 2. Distribution of cell diameters of small spherical unicellular morphotypes

(frequency is expressed as the ratio between the number of cells of a given dia-
meter and the total number of measured cells).

Colonies mucilaginous, sheath colorless sometimes yellowish with dis-

tinct margins often diffluent. Young colonies more or less spherical, later
irregular, sometimes lobate. Cells irregularly and densely arranged within
the colonies, blue-green, spherical, 0.82 ± 0.04 (0.76–0.86) µm in diameter.
Fig. 6a–c.
O c c u r r e n c e : BSU, L10, LBU.
M o r p h o t y p e : U06
Colonies mucilaginous, sheath colorless with indistinct margin. Cells ir-
regularly and more or less densely arranged within the colonies; cells blue-
green, spherical, 1.13 ± 0.10 (0.94–1.39) µm in diameter. Fig. 5a–c.
O c c u r r e n c e : L73, L68, L1, L52, LMA, L8, L63, L23, L74.
C o m m e n t s : We have distinguished morphotypes U05 and U06 mainly

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 Cyanobacteria in mats of East Antarctic lakes 181

Fig. 3. Cell widths and lengths of small oval or cylindrical unicellular morphotypes.

on the basis of the cell diameter (Fig. 3). Both correspond to the morpho-
logical description of Aphanocapsa cf. holsatica, which is a common cosmo-
politan planktonic taxon from freshwater habitats and therefore its ecology
does not correspond to the biotope of our specimens.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : This species is re-
ported for the first time from Antarctica.

Aphanocapsa cf. hyalina (LYNGBYE) HANSGIRG 1892

M o r p h o t y p e : U09
Colonies more or less spherical or irregularly spherical. Mucilage ge-
latinous, colorless to yellowish with distinct margins; cells irregularly and
loosely arranged within the colonies, blue-green, spherical, 2.42 ± 0.25
(1.84–2.80) µm in diameter. Fig. 5h–i.
O c c u r r e n c e : L53b, L65, L52b, LG2, L59b.

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 182 A. TATON et al.

Fig. 4. Distribution of cell widths and lengths of small oval or cylindrical unicel-
lular morphotypes (frequencies are expressed as the ratio between the number of
cells of a given width or length and the total number of measured cells).

C o m m e n t s : This is a freshwater benthic, epipelic taxon, mainly found in

mountains of central Europe. However, it was also reported from Africa in
clear swamps, peaty waters, and small stagnant water bodies.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : This species is re-
ported for the first time from Antarctica.

Aphanothece cf. elabens (BRÉBISSON) ELENKIN 1938

B a s i o n y m : Micraloa elabens BRÉBISSON 1842

S y n o n y m : Microcystis elabens (BRÉBISSON) KÜTZING 1846; Polycystis
elabens (BRÉBISSON) KÜTZING 1849; Microcystis elabens var. maior BACH-
MANN 1921; Microcystis aphanothecoïdes ZALESSKY 1926; Aphanothece ela-
bens var. minor NYGAARD 1929; Coccochloris elabens (BRÉBISSON) DROUET
ET DAILY 1956
M o r p h o t y p e : U03
Colonies mucilaginous, with homogeneous, colorless envelopes, more or
less delimited to diffluent. Cells pale blue-green, oval to cylindrical, 2.00 ±
0.24 (1.53–2.45) µm wide, 3.24 ± 0.51 (2.26–4.31) µm long. Fig. 6c–e.
O c c u r r e n c e : L43, L51, L70, L59, L13, L23, L67, L74.
C o m m e n t s : This is a freshwater, epipelic and metaphytic taxon, some-
times free-floating in lakes recorded from temperate and tropical regions.

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 Cyanobacteria in mats of East Antarctic lakes 183

Fig. 5. Photomicrographs of the small spherical Chroococcales: (a–c) Aphano-

capsa cf. holsatica; (d) cf. Chamaesiphon; (e–g) Chlorogloea sp.; (h–i) Aphanocapsa
cf. hyalina [scale bars = 10 µm].

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 184 A. TATON et al.

E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : This species is re-

ported for the first time from Antarctica.

Aphanothece cf. microscopica NÄGELI 1849

M o r p h o t y p e : U04
Colonies mucilaginous, with homogeneous, more or less delimited to dif-
fluent colorless envelopes. Cells more or less densely arranged within the
colonies, sometimes with their own colorless simple envelope, pale blue-
green to olive green, widely oval to cylindrical, 2.59 ± 0.25 (2.19–3.46) µm
wide, 3.94 ± 0.47 (3.15–5.52) µm long. Fig. 6f–g.
O c c u r r e n c e : L64, L43, L65, L51, L13, L23, L67.
C o m m e n t s : This taxon is usually found in the benthic and epiphytic
communities of freshwater environments in temperate regions.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Palmer Peninsula
(PEN) (PRESCOTT 1979).

Aphanothece sp.

M o r p h o t y p e : U01
Colonies mucilaginous, with homogeneous, usually diffluent, rarely dis-
tinctly delimited envelopes. Cells pale blue-green, oval to cylindrical, 1.06 ±
0.13 (0.84–1.38) µm wide, 2.44 ± 0.41 (1.54–3.31) µm long. Fig. 6a–b.
O c c u r r e n c e : L64, LFO, L53b, L1, L52b, LPS, L66, LG1.

Asterocapsa sp. (Gloeocapsa cf. rupestris KÜTZING 1845)

M o r p h o t y p e : ASru
Cells solitary, in groups of a few cells or in more or less spherical mi-
croscopic colonies. Cells or groups of cells enveloped by distinct, delimited,
firm, lamellate or not, yellow to yellow-brown sheath (with a granular or
spiny surface). Subcolonies with colored sheaths are sometimes surrounded
by wide, more or less spherical, colorless, homogeneous and well delimited
envelopes. Subcolonies sometimes confluent in the centre of large colonies.
Cells pale blue-green, subspherical to polygonal, 5.91 ± 1.32 (4.32–10.64)
µm wide, 7.75 ± 1.55 (4.80–10.76) µm long. Fig. 7a–d.
O c c u r r e n c e : L52b, LG2.
C o m m e n t s : This is a subaerophytic taxon found on periodically wet
rocks, from lowlands to high mountains and is widely distributed.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : This species is re-
corded for the first time in Antarctica.

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 Cyanobacteria in mats of East Antarctic lakes 185

Fig. 6. Photomicrographs of the small oval or cylindrical Chroococcales: (a–b)

Aphanothece sp.; (c–e) Aphanothece cf. elabens; (f–g) Aphanothece cf. microscop-
ica; (h) Rhabdonema sp. [scale bars = 10 µm].

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 186 A. TATON et al.

Chamaesiphon subglobosus (ROSTAFINSKI) LEMMERMANN 1910

B a s i o n y m : Sphaerogonium subglobosum ROSTAFINSKI 1883

M o r p h o t y p e : CMsu
Cells solitary or aggregated in small irregular colonies. Cell division
asymmetrical. Cells heteropolar, attached or not by their bases to the
sheath of other cells of the colony (or to the substrate), pale blue-green,
sometimes slightly yellowish. Mother cells spherical to ovoid, 3.79 ± 0.66
(2.40–4.92) µm wide, 4.38 ± 0.89 (2.40–5.84) µm long. Sheath pseudovaginae
colorless, not clearly distinct or indistinct. Reproduction by a single exocyte
differentiating at the cell apex; exocyte spherical, 2.31 ± 0.28 (1.48–2.80) µm
in diameter. Fig. 7e.
O c c u r r e n c e : LG1, L13.
C o m m e n t s : This taxon is usually found in the epilithic and epiphytic
communities of freshwater lakes and streams, from lowlands to mountains,
mainly in central Europe.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Periphyton of lakes
of Hope Bay (PEN) (VINOCUR & PIZARRO 1995), streams of Dry Valleys
(SVL) (BROADY 1982), plankton of lakes and ponds of Hope Bay (PEN)
(TELL et al. 1995), plankton, felt and ice bubble of Lake Otero, Cierva Point,
Hope Bay (PEN) (MATALONI et al. 1998).

Cf. Chamaesiphon sp.

M o r p h o t y p e : U08
Colonies more or less mucilaginous with cells arranged irregularly. Cells
with irregular shape, from spherical to pear-shaped. Cell diameter variable
in the same colonies, 2.22 ± 0.45 (1.51–2.98) µm in diameter. Fig. 5d.
O c c u r r e n c e : LPS, L63, L71, and L74.
C o m m e n t s : This taxon probably belongs to the genus Chamaesiphon
and the variability of the cell diameter within the same colony is due to
asymmetrical division. However, this morphotype was less typical for the
genus in comparison to Chamaesiphon cf. subglobosus described above.

Chondrocystis cf. dermochroa (NÄGELI) KOMÁREK ET ANAGNOSTIDIS 1995

B a s i o n y m : Gloeocapsa dermochroa NÄGELI 1849

M o r p h o t y p e : CHde
Colonies composed of densely packed subcolonies. Subcolonies conflu-
ent in the center of large colonies, spherical, subspherical to polygonal. Sub-
colonies enveloped by a more or less firm to diffluent, slightly or not lamel-
late, yellow to yellow-brown sheath. Cells blue-green, spherical, 2.86 ± 0.64
(2.08–4.80) µm in diameter. Fig. 7f–h.
O c c u r r e n c e : L64, LGR, LSP, L53b, L65, L52b, L52, LPS, L70, LMA,
LG2, L66, L59b, LSN, L61, L72.

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 Cyanobacteria in mats of East Antarctic lakes 187

Fig. 7. Photomicrographs of the Chroococcales: (a–d) Asterocapsa sp.; (e) Chamae-

siphon subglobosus; (f–h) Chondrocystis cf. dermochroa. [scale bars =10 µm].

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 188 A. TATON et al.

C o m m e n t s : This is a subaerophytic taxon usually found on rocks with

periodically flowing water and in the splash zone of the lake littorals in cen-
tral Europe.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Mats of lakes and
ponds of Inexpressible Island (NVL) (FUMANTI et al. 1997); plankton of
lakes and ponds of Hope Bay (PEN) (IZAGUIRRE et al. 1993, TELL et al.
1995, IZAGUIRRE et al. 1998) and soils of Mt McGee (NVL) (CAVACINI
2001).

Chlorogloea sp.

M o r p h o t y p e : U07
Colonies mucilaginous, sheath colorless with indistinct margin. Cells ir-
regularly, loosely or densely arranged in a common mucilage, spherical, 2.13
± 0.36 (1.34–3.61) µm in diameter. Fig. 5e–g.
O c c u r r e n c e : L49, L64, LFO, LGR, L68, L53b, L42, L65, L1, L52b, L52,
L18, LJA, BBA, BSU, LPS, L70, LMA, LG2, L66, LG1, L61, L72, L34, L14,
L10, L8, L71, L13, LBU, L23, L67, L74.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Several species of
this genus, C. fritschii MITRA, C. microcystoides GEITLER, and C. purpurea
GEITLER were found in the plankton or the periphyton of lakes and ponds
of Hope Bay (PEN) (IZAGUIRRE et al. 1993, 1998, TELL et al. 1995, VINOCUR
& PIZARRO 1995) and Chlorogloea sp. in streams of King Georges Island
(South Shetland Islands, MAR) (KOMÁREK & KOMÁREK 2001).

Gloeocapsa cf. alpina (NÄGELI) BRAND 1900

S y n o n y m : Gloeocapsa ambigua NÄGELI in KÜTZING 1849; G. fuscolutea

KIRCHNER 1878
M o r p h o t y p e : GLal
Colonies composed of subcolonies, with wide mucilaginous envelopes,
which are granulated, finely lamellate, blue to dark violet; subcolonies with
a colored sheath sometimes surrounded by a wide, more or less spherical,
colorless, homogeneous and well delimited envelope. Cells blue-green,
spherical, 4.83 ± 0.38 (3.60–5.60) µm wide, 5.18 ± 0.35 (4.52–6.40) µm long.
Fig. 8a.
O c c u r r e n c e : LG2.
C o m m e n t s : This is a subaerophytic taxon found on rocks in mountains
and which is widely distributed all over the world.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Periphyton of Lake
Encantado of Hope Bay (PEN) (VINOCUR & PIZARRO 1995), streams of
Dry valleys (SVL) (BROADY 1982), edaphic, epilithic and epiphytic com-
munities of terrestrial environments in the Vestfold Hills (PEL) (BROADY
1986), epilithic and epiphytic community of terrestrial environments of Ed-
ward VII Peninsula (MBL) (BROADY 1989)

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 Cyanobacteria in mats of East Antarctic lakes 189

Fig. 8. Photomicrographs of the Chroococcales: (a) Gloeocapsa cf. alpina; (b–d)

Gloeocapsa cf. compacta; (e–f) Gloeocapsa cf. sanguinea. [scale bars = 10 µm].

Gloeocapsa cf. compacta KÜTZING 1845

M o r p h o t y p e : GLco
Colonies composed of tightly packed subcolonies. Subcolonies some-
times solitary, ovoid or irregularly ovoid. Mucilaginous envelopes colorless
to brown-violet or black. Cells blue-green, spherical, 3.39 ± 0.40 (2.56–4.40)
µm in diameter. Fig. 8b–d.
O c c u r r e n c e : LSP, L53b, LG2, L10.
C o m m e n t s : Specimens slightly bigger than the original description. Sub-
aerophytic taxa found from lowlands to mountains of central Europe, com-
mon in high mountains.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : This species is re-
ported for the first time in Antarctica.

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 190 A. TATON et al.

Gloeocapsa cf. sanguinea (AGARDH) KÜTZING 1843

S y n o n y m : Gloeocapsa itzigsohnii BORNET in ZOPF 1882

M o r p h o t y p e : GLsa
Colonies composed of subcolonies. Subcolonies enveloped by a firm,
later diffluent, not or slightly lamellate sheath, intensely red near the cells,
with outer envelope slightly reddish to colorless. Subcolonies with a colored
sheath sometimes surrounded by a wide, more or less spherical, colorless,
homogeneous and well delimited envelope. Cells pale blue-green, spheri-
cal, 4.76 ± 0.46 (3.76–5.40) µm in diameter. Fig. 8e–f.
O c c u r r e n c e : LG2.
C o m m e n t s : This is a subaerophytic taxon found on wet rocks in high
mountains. It is possibly cosmopolitan and is also found in Northern re-
gions up to subpolar areas.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Terrestrial environ-
ments of King George Island (South Shetland Islands, MAR) (KOMÁREK
1999).

Rhabdonema sp.

M o r p h o t y p e : U02
Colonies mucilaginous, with colorless and diffluent envelope. Cells pale
blue-green, cylindrical, straight or slightly arched, 1.05 ± 0.14 (0.81–1.29)
µm wide, 4.01 ± 0.72 (2.69–5.42) µm long. Fig. 6h.
O c c u r r e n c e : L64, L65, L52b.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : This genus is re-
ported for the first time in Antarctica.

Oscillatoriales

The thin Oscillatoriales were very difficult to identify in the field samples.
Therefore, for each sample, pictures of each cyanobacterium appearing
to be different in light microscopy were taken. Based on the pictures, cell
measurements were realized (Fig. 9) and morphological diacritical traits
determined to distinguish the different morphotypes.

Leptolyngbya sp.

M o r p h o t y p e : LPP02
Filaments straight or curved. Trichomes sometimes with a very thin
sheath, slightly or not constricted at the cross-wall. Cells blue-green, cylin-
drical, longer than wide, 0.89 ± 0.14 (0.61–1.12) µm wide, 4.72 ± 1.65 (1.81–
10.23) µm long, sometimes with a granule near the cross-wall. End cells
rounded. Fig. 10a–b

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 Cyanobacteria in mats of East Antarctic lakes 191

Fig. 9. Distribution of cell widths and lengths of thin Oscillatoriales (frequencies

are expressed as the ratio between the number of cells of a given width or length
and the total number of measured cells).

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 192 A. TATON et al.

O c c u r r e n c e : L44, L43, L65, L51, L1, LJA, BBA, LPS, L70, LMA, LG2,
L59b, LG1, L61, L72, L59, L63, L71, L13, L67.
C o m m e n t s : This morphotype is characterized by a wide variation of the
cell length (Fig. 10). It corresponds to several species including, for exam-
ple, Leptolyngbya antarctica (W. et G. S. WEST) ANAGNOSTIDIS et KOMÁREK
1988 (Bas: Phormidium antarcticum W. et G. S. WEST 1911), a possible en-
demic species previously found in streams, lakes and ponds of continental
and maritime Antarctica.

Phormidium autumnale (AGARDH) TREVISAN ex GOMONT 1892

S y n o n y m : Phormidium membranaceum KÜTZING ex GOMONT 1892;

Phormidium autumnale var. minus GARDNER 1927; Lyngbya autumnalis
(AGARDH) ex BOURRELLY 1970; Lyngbya antliaria (MERTENS in JÜRGENS)
HANSGIRG ex HANSGIRG 1892; Phormidium pannosum KÜTZING ex GO-
MONT 1892
M o r p h o t y p e : OSs1
Filaments straight. Trichomes not constricted or very slightly constricted
at the cross-wall, usually attenuated and sometime slightly hooked at the
apex. Cells blue-green, cylindrical, shorter than wide, 7.11 ± 0.47 (6.16–7.84)
µm wide, 3.21 ± 0.58 (2.04–4.56) µm long. Terminal cells more or less capi-
tate, sometimes with a calyptra. Fig. 10c.
O c c u r r e n c e : LFO, L52.
C o m m e n t s : This taxon is frequently described from Antarctica, but it
comprises different morphotypes and ecotypes from which several indi-
vidual species were described (KOMÁREK 1999). In the “Phormidium au-
tumnale group”, KOMÁREK (1999) distinguished three species found in dif-
ferent biotopes of King George Island. Our specimens correspond to the
morphological description of Phormidium attenuatum but do not possess
the same ecological specificities.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Mats in lakes and
ponds of the Ross Island (RSR) (BROADY & KIBBLEWHITE 1991) and the
McMurdo Ice Shelf (RSR) (BROADY & KIBBLEWHITE 1991, NADEAU et
al. 2001); plankton in lakes and ponds of Hope Bay (PEN) (IZAGUIRRE et
al. 1998, MATALONI et al. 1998) and McMurdo Ice Shelf (RSR) (JAMES et
al. 1995); streams in Dry Valleys (SVL) (MCKNIGHT et al. 1998); sublithic,
edaphic, epiphytic communities of terrestrial environments of Edward VII
Peninsula (MBL) (BROADY 1989); soils of Cierva Point, Hope Bay (PEN)
(MATALONI et al. 2000) and Ongul Island (“EL”) (OHTANI et al. 2000);
sublithic, edaphic, epiphytic communities of terrestrial environments in
the Vestfold Hills (PEL) (BROADY 1986); soils and pond sediments on La
Gorce Mountains (QML) (BROADY & WEINSTEIN 1998); ice and snow at
Cierva Point, Hope Bay (PEN) (MATALONI & TESOLIN 1997); different bio-
topes in continental and maritime Antarctica (PRESCOTT 1979).

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 Cyanobacteria in mats of East Antarctic lakes 193

Fig. 10. Photomicrographs of the Oscillatoriales: (a–b) Leptolynbya sp.; (c) Phor-
midium autumnale; (d) Phormidium murrayi; (e–f) Phormidium priestleyi; (g–h)
Pseudanabaena frigida; (i) Pseudanabaena cf. amphigranulata; (j) Pseudanabaena
cf. tremula; (k) Schizothrix lacustris; (l) Schizothrix cf. simplicior; (m) Schizothrix cf.
tenuis; (n) cf. Jaaginema sp. [scale bars = 2 µm].

Phormidium murrayi (W. et G. S. WEST) ANAGNOSTIDIS et KOMÁREK 1988

B a s i o n y m : Lyngbya murrayi W. et G. S. WEST 1911

M o r p h o t y p e : LYs1
Filaments straight, 3.57 ± 0.57 (2.52–5.00) µm wide. Trichomes en-
sheathed, not constricted or very slightly constricted at the cross-walls,
sometimes slightly attenuated at the apex. Cells blue-green, cylindrical,
shorter than wide, 2.88 ± 0.28 (2.44–3.52) µm wide, 5.63 ± 0.92 (4.08–8.00)
µm long. Terminal cell rounded. Fig. 10d.
O c c u r r e n c e : L52, BSU, L70, LSN.
C o m m e n t s : This taxon is found in stagnant, mostly salty waters, in Ant-
arctica and possibly also in the Arctic regions.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Mats in ponds of
the McMurdo Ice shelf (RSR) (NADEAU et al. 2001), Green Lake on Ross

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 194 A. TATON et al.

Island (RSR) (WEST & WEST 1911); cryoconites on glaciers in Dry Valleys
(SVL) (BROADY & KIBBLEWHITE 1991); epiphytic of terrestrial environ-
ments of Vestfold Hills (PEL) (BROADY et al. 1984); Ross Island (RSR)
(PRESCOTT 1979).

Phormidium priestleyi FRITSCH 1917

S y n o n y m : Lyngbya priestleyi (FRITSCH) SENNA 1983

M o r p h o t y p e : LPP01
Filaments straight and curved. Trichomes ensheathed, constricted at the
cross-wall. Sheath hyaline. Cells blue-green, barrel-shaped, shorter than
wide to slightly longer than wide, 1.13 ± 0.15 (0.88–1.50) µm wide, 1.42 ±
0.25 (0.88–1.93) µm long. End cells rounded. Fig. 10e.
M o r p h o t y p e : LPP06
O c c u r r e n c e : L49, LFO, L52b, L18.
Filaments straight and curved. Trichomes ensheathed, constricted at the
cross-wall. Sheath hyaline, sometimes blackish. Cells blue-green, barrel-
shaped, quadratic to longer than wide, 1.75 ± 0.17 (1.45–2.18) µm wide, 2.97
± 0.71 (1.99–5.05) µm long. End cells rounded. Fig. 10f.
O c c u r r e n c e : L68, L18, BSU, L66, L63, L74.
C o m m e n t s : On the basis of the cell size (Fig. 9), we have distinguished
two morphotypes that correspond to the description of Phormidium priest-
leyi. This species is characteristic of the epilithic communities of flowing
and rarely stagnant waters in Maritime Antarctica and is a possible Antarc-
tic endemic species (KOMÁREK 1999, KOMÁREK & ANAGNOSTIDIS 2005).
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Lakes and ponds of
the Dry Valleys (SVL) (PRESCOTT 1979, BROADY et al. 1984); periphyton of
lakes and ponds of Hope Bay (PEN) (VINOCUR & PIZARRO 1995); plank-
ton, felt and ice bubble of Lake Otero at the Cierva Point, Hope Bay (PEN)
(MATALONI et al. 1998); streams of King Georges Island (South Shetland
Islands, MAR) (KOMÁREK 1999, KOMÁREK & KOMÁREK 2001); soils of sev-
eral locations (NVL) (CAVACINI 2001); the chasmoendolithic community of
terrestrial environments at Mawson Rock (MRL) (BROADY et al. 1984).

Pseudanabaena frigida (FRITSCH) ANAGNOSTIDIS 2001

B a s i o n y m : Phormidium frigidum FRITSCH 1912

S y n o n y m : Leptolyngbya frigida (FRITSCH) ANAGNOSTIDIS et KOMÁREK
1988 p.p.
M o r p h o t y p e : LPP05
Filaments straight and curved. Trichomes slightly constricted at the cross-
walls. Cells blue-green, cylindrical, longer than wide, 1.32 ± 0.24 (0.77–1.73)
µm wide, 2.76 ± 0.74 (1.78–4.76) µm long. End cells rounded or conical.
Fig. 10g.
O c c u r r e n c e : L44, L42, L51, LJA, LPS, L10, L23.

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 Cyanobacteria in mats of East Antarctic lakes 195

M o r p h o t y p e : LPP07
Filaments straight and curved. Trichomes ensheathed, slightly con-
stricted at the cross-walls, sometimes apparently with a space between the
cells. Sheath hyaline. Cells blue-green, cylindrical, quadratic to longer than
wide, 1.34 ± 0.13 (1.10–1.61) µm wide, 2.17 ± 0.57 (1.30–3.43) µm long. End
cells rounded. Fig. 10h.
O c c u r r e n c e : L8, LBU, L23.
C o m m e n t s : We have distinguished two morphotypes LPP05 and LPP07.
Cells are always longer than wide for LPP05 whereas LPP07 sometimes
possesses quadratic cells. In addition, the trichomes of LPP07 are usually
ensheathed, whereas LPP05 does not have a sheath. Both characters are
very plastic and both morphotypes correspond to the description of Pseu-
danabaena frigida. This species is usually found in stagnant and flowing,
freshwater environments of Antarctica, Arctica and North of Europe.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Mats and plankton
of lakes and ponds in the Dry Valleys (SVL) (BROADY 1982, WHARTON et
al. 1983, PARKER & WHARTON 1985, SIMMONS et al. 1993a, SPAULDING et al.
1994, MCKNIGHT et al. 2000); mats of lakes on Inexpressible Island (NVL)
(FUMANTI et al. 1997) of freshwater lakes on the Schirmacher Oasis (QML)
(KOMÁREK & RUCZICKA 1966); plankton of lakes and ponds of Hope Bay
(PEN) (IZAGUIRRE et al. 1998); streams in the Dry Valleys (SVL) (BROADY
et al. 1984); sublithic, epiphytic and edaphic communities of terrestrial envi-
ronments in the Vestfold Hills (PEL) (BROADY 1986), soils of Cierva Point,
Hope Bay (PEN) (MATALONI et al. 2000); soils of several locations (NVL)
(CAVACINI 2001); ice and/or snow in McMurdo Sound (RSR) (FRITSCH 1912
according to PRESCOTT 1979) and Cierva Point, Hope Bay (PEN) (MATAL-
ONI & TESOLIN 1997); plankton, felt and ice bubble of Lake Otero at Cierva
Point, Hope Bay (PEN) (MATALONI et al. 1998), freshwater environment
of King George Island (South Shetland, MAR) (KOMÁREK 1999), Folmar
Island (PRESCOTT 1979).

Pseudanabaena cf. amphigranulata (VAN GOOR) ANAGNOSTIDIS 2001

B a s i o n y m : Oscillatoria amphigranulata VAN GOOR 1918

S y n o n y m : Limnothrix amphigranulata (VAN GOOR) MEFFERT 1988 sensu
VAN GOOR; Pseudanabaena galeata BÖCHER f. tenuis (BÖCHER) POLJANSKIJ
in HOLLERBACH et al. 1953
M o r p h o t y p e : LPP08
Filaments straight and curved. Trichomes ensheathed slightly constricted
at the cross-walls. Cells blue-green, cylindrical, longer than wide, 1.70 ± 0.12
(1.56–1.90) µm wide, 5.35 ± 0.76 (3.92–6.10) µm long, with a granule or a gas
vesicle near the cross-wall. Terminal cell rounded. Fig. 10i.
O c c u r r e n c e : L18, L74.
C o m m e n t s : There are various ecological concepts for this species
(KOMÁREK 1999). It was found in benthic, metaphytic communities or in

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 196 A. TATON et al.

periphyton of freshwater environments or in mud of lakes and various wa-

ter bodies containing H2S. Possibly widely distributed over the world.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Plankton of Prasiola
stream of Hope Bay (PEN) (VINOCUR & PIZARRO 1995).

Pseudanabaena cf. tremula CASAMATTA et JOHANSEN 2005

M o r p h o t y p e : LPP03
Filaments straight and curved. Trichomes ensheathed, not to slightly con-
stricted at the cross-walls, sometimes apparently with a space between the
cells. Cells blue-green, cylindrical, longer than wide, 1.20 ± 0.12 (0.94–1.38)
µm wide, 3.77 ± 1.18 (2.50–6.85) µm long, sometimes with a granule or a gas
vesicle near the cross-wall. End cells rounded. Fig. 10j.
O c c u r r e n c e : L49, L68, L66, L63.
C o m m e n t s : This species was recently described by CASAMATTA et al.
(2005) on the basis of the strain UTCC471 isolated from the benthic sub-
strate in a pond on Bylot Island, Northwest Territories, Canada. However,
we were unable to observe the characteristic trembling motility of this
taxon since our samples were fixed.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : This species is re-
ported for the first time in Antarctica.

Schizothrix lacustris A. BRAUN ex GOMONT 1892

S y n o n y m : Microcoleus braunii H ANSGIRG 1892, Inacystis lacustris

(GOMONT) FORTI 1907, Schizothrix lacustris var. caespitosa GOMONT 1892,
Schizothrix fasciculata GOMONT sensu KANN 1978
M o r p h o t y p e : SCs1
Several trichomes within a common sheath. Trichomes slightly con-
stricted at the cross-walls. Sheath colorless. Cells blue-green to pinkish,
cylindrical, more than two times longer than wide in average, 1.28 ± 0.23
(1.12–2.40) µm wide, 3.41 ± 0.61 (2.40–4.84) µm long. End cells conical.
Fig. 10k.
O c c u r r e n c e : L49, L64, LFO, LGR, L51, L1, L52b, LG2, L59b, LSN.
C o m m e n t s : This species was found in freshwater lakes and streams,
mainly of mountains in Europe, North America and Asia.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Melt water at Marble
point on Ross Island (RSR) (PRESCOTT 1979); Mt. Suess (SVL) (PRESCOTT
1979).

Schizothrix cf. simplicior SKUJA 1964

M o r p h o t y p e : SCs3
Several trichomes within a common sheath. Trichomes constricted at the
cross-walls. Sheath colorless to violet or blackish. Cells blue-green, barrel-

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 Cyanobacteria in mats of East Antarctic lakes 197

Fig. 11. Distribution of cell widths and lengths of Schizothrix morphotypes (fre-
quencies are expressed as the ratio between the number of cells of a given width or
length and the total number of measured cells).

shaped, 1.75 ± 0.17 (1.36–2.08) µm wide, 2.18 ± 0.58 (1.00–3.68) µm long.

End cells rounded. Fig. 10l.
O c c u r r e n c e : L68, L53b, L65, L52, LPS, LMA, LG2, L66, LSN, L61, L10,
L63, L74.
C o m m e n t s : This species was found in freshwater alpine and subalpine
ponds and in glacier lakes in Austria and North Sweden. The cell dimen-
sions of two populations of this morphotype, one with colorless sheath and
the other with blackish sheath, were compared but did not reveal any dif-
ferences (Fig. 11).
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : This species is re-
ported for the first time in Antarctica.

Schizothrix cf. tenuis VORONICHIN 1923

M o r p h o t y p e : SCs2
Several trichomes within a common sheath. Trichomes slightly con-
stricted at the cross-walls. Sheath colorless. Cells blue-green, barrel-shaped
to cylindrical, 1.43 ± 0.11 (1.12–1.68) µm wide, 2.17 ± 0.41 (1.44–3.20) µm
long. End cells rounded to conical. Fig. 10m.
O c c u r r e n c e : L49, LSP.
C o m m e n t s : This taxon is known from springs and streams.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : This species is re-
corded for the first time in Antarctica.

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 198 A. TATON et al.

Cf. Jaaginema sp.

M o r p h o t y p e : LPP04
Filaments straight and curved. Trichomes not constricted at the cross-
wall. Cells blue-green, cylindrical, longer than wide, 1.39 ± 0.10 (1.18–1.51)
µm wide, 4.68 ± 0.62 (4.13–5.99) µm long. End cells rounded. Fig. 10n.
O c c u r r e n c e : L68.
C o m m e n t s : This morphotype was rarely observed in our samples.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : This genus is re-
ported for the first time in Antarctica.

Nostocales
Calothrix cf. parietina (NÄGELI) THURET ex BORNET et FLAHAUT 1886

M o r p h o t y p e : CAs1
Filaments unbranched, 12.20 ± 1.35 (7.64–14.24) µm wide at the base in
the largest part of the filament. Trichomes tapering with a long colorless
hair. Sheath often yellow-brown and layered. Cells blue-green, 7.08 ± 0.63
(5.44–8.20) µm wide and 4.14 ± 0.97 (2.88–6.72) µm long in the largest part
of the trichome. Heterocyst basal, 7.22 ± 1.00 (4.00–8.56) µm wide, 5.00 ±
0.68 (4.00–6.96) µm long. Fig. 12a–b.
O c c u r r e n c e : L68, L52, LMA, LG2, L61, L10, L63.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Mats of lakes on
Inexpressible Island (NVL) (FUMANTI et al. 1997); streams in Dry Val-
leys (SVL) (BROADY 1982); epilithic, chasmoendolithic, edaphic, epiphytic
communities of terrestrial environments of Vestfold Hills (PEL) (BROADY
1986); soils of several locations (NVL) (CAVACINI 2001); Ross Island (RSR),
Walcott Glacier and Budd Coast (WL) (PRESCOTT 1979).

Coleodesmium cf. scottianum WELSH 1965

M o r p h o t y p e : COsc
Filaments falsely branched, forming fascicles, (polarized); one or sev-
eral trichomes in the same sheath; 13.44 ± 4.52 (7.01–22.32) µm wide if
there is only one trichome in the sheath; sheaths firm, slightly lamellate,
opened at the apex, yellow-brown. Trichomes polarized with basal hetero-
cysts, constricted at the cross walls; false branching often initiating at the
intercalary heterocysts; trichomes sometimes slightly narrower at the end.
Cells blue-green, shortly barrel-shaped, 9.32 ± 1.02 (7.48–11.24) µm wide,
5.78 ± 1.46 (3.56–9.00) µm long; end cells rounded. Heterocysts 10.35 ± 1.93
(6.56–13.16) µm wide, 13.41 ± 3.98 (8.92–22.24) µm long. Akinetes absent.
Fig. 12d–f.
O c c u r r e n c e : LJA, LSN, L61.
C o m m e n t s : Our specimens morphologically, but not ecologically, corre-
spond to two species reviewed by KOMÁREK & WATANABE (1990), C. scot-

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 Cyanobacteria in mats of East Antarctic lakes 199

Fig. 12. Photomicrographs of the Nostocales: (a–b) Calothrix sp.; (c) Dichothrix
sp.; (d–f) Coleodesmium cf. scottianum; (g) Petalonema cf. involvens; (h–j) Nostoc
sp. [scale bars = 10 µm].

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 200 A. TATON et al.

tianum WELSH 1965 described from cataracts in streams of Okavango (S. W.

Africa) and C. swazilandicum WELSH 1965 from stones in fast running
streams of Swaziland (S. Africa).
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Until now, the genus
Coleodesmium (Desmonema wrangelii (AGARDH) ex BORN. et FLAH.) was
only found in MAR (PRESCOTT 1979).

Dichothrix sp.

M o r p h o t y p e : DIsp
Filaments often falsely branched, 11.14 ± 2.56 (7.48–17.56) µm wide at
the base in the largest part of the filaments. Trichomes tapering at the apex
but without a long terminal hair. Sheath often yellow-brown and stratified.
Cells blue-green, 6.53 ± 2.10 (0.69–11.72) µm wide and 4.30 ± 1.01 (3.04–
7.20) µm long in the largest part of the trichome. Heterocyst basal, 6.98 ±
1.78 (4.00–12.40) µm wide, 6.70 ± 1.14 (4.88–9.40) µm long. Fig. 12c.
O c c u r r e n c e : LFO, L53b, L52b, L52, LG2.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Until now, this genus
was only found in Maritime Antarctica (e. g. HIRANO 1965, THÉRÉZIEN 1965
according to PRESCOTT 1979).

Nostoc sp.

M o r p h o t y p e : NOsp
Thallus microscopical, gelatinous, amorph or spherical, later irregularly
spherical, lobate. Filaments within colony irregularly coiled and loosely or
densely agglomerated; sheath around trichomes sometimes present, colo-
nial mucilage sometimes yellowish-brown. Cells almost spherical, 4.45 ±
0.77 (3.20–6.36) µm wide, 4.37 ± 0.91 (2.16–7.20) µm long. Heterocysts soli-
tary, intercalary, 4.97 ± 0.94 (3.76–7.72) µm wide, 4.86 ± 1.24 (2.84–7.28) µm
long. Fig. 12h–j.
O c c u r r e n c e : L64, L64b, L43, LFO, LGR, L68, LSP, L53b, L42, L65, L51,
L1, L52b, L52, L18, LJA, BSU, LMA, LG2, L66, L59b, LG1, L72, L14, L8,
L59, L63, L71, L13, LBU.
C o m m e n t s : There are important variations in cell dimensions. However,
no gaps were found and there is a continuum from smaller cells to larger
cells.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Several species
of this genus were frequently found in diverse terrestrial, lotic and lenthic
biotopes of continental and maritime Antarctica. N. antarcticum W. WEST et
G. S. WEST, N. commune VAUCH. ex BORN. et FLAH., N. disciforme FRITSCH,
N. fuscescens FRITSCH, N. longstaffii FRITSCH, N. microscopicum CARM. ex
BORN. et FLAH., N. punctiforme (KÜTZ.) ex HARIOT. N. sphaericum VAUCH.
ex BORN. et FLAH. was found in the microbial mats of lakes and ponds of Ross
Sea Region (RSR) and/or the Dry Valleys (SVL) (e. g. FRITSCH 1912, WHAR-

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 Cyanobacteria in mats of East Antarctic lakes 201

TON et al. 1983, PARKER & WHARTON 1985, SIMMONS et al. 1993). N. commune
was also found on Inexpressible Island (NVL) (FUMANTI et al. 1997).

Petalonema cf. involvens (A. BRAUN) ex MIGULA

B a s i o n y m : Symphyosiphon involvens A. BRAUN 1856

S y n o n y m : Scytonema involvens RABENH. ex BORN. et FLAH. 1886
M o r p h o t y p e : PEin
Filaments falsely branched, 20.52 ± 3.73 (13.84–28.96) µm wide. Sheath
firm, very thick, distinctly funnel-like, lamellate and yellow-brown. Tri-
chomes constricted at the cross-wall; cells barrel-shaped, shorter in the ter-
minal parts than in the median parts; apical cells rounded; cells 6.51 ± 0.99
(4.52–8.52) µm wide, 5.93 ± 1.17 (4.00–8.52) µm long. Heterocysts interca-
lary, more or less spherical and usually wider than vegetative cells, 8.18 ±
1.16 (6.64–11.12) µm wide and 9.00 ± 1.88 (5.40–13.12) µm long. Akinetes
missing. Fig. 12g.
O c c u r r e n c e : L64, LG2.
C o m m e n t s : Lakes in Germany, California, Australia.
E c o l o g y a n d d i s t r i b u t i o n i n A n t a r c t i c a : Until now, Petalo-
nema densum (A. BRAUN) MIGULA was the only species of this genus found
in Antarctica: Kerguelen Island (MAR) (PRESCOTT 1979).

Diversity, ecology and biogeography

The studied samples contained 30 taxa corresponding to 33 morphotypes
belonging to 13 genera of Chroococcales, Oscillatoriales and Nostocales.
No representative of the order Stigonematales was recorded.
A semi-quantitative estimation of the abundance of the different
morphotypes in the samples was realized (Fig. 13). The presence/absence
data were included in a 56–lake dataset representing the limnological
diversity among more than 150 lakes and ponds in the Larsemann Hills
and Bølingen Regions to identify the abiotic conditions associated with
cyanobacterial and diatom communities (SABBE et al. 2004). These authors
distinguished 5 mat physiognomies and showed that the mat physiognomy
in the studied lakes was strongly correlated to lake depth but not to the
salinity: finely laminated prostrate mats were restricted to deeper lakes (>
7 m); less structured prostrate mats occurred between 2 and 16 m; flake
mats, consisting of small (1–2 cm) plate-like growths of cyanobacteria,
were confined to shallow lakes up to 2.3 m deep; epipsammic (interstitial
and epilithic) mats consisted of an organic matrix with a high amount of
embedded inorganic sediment; the latter were observed in shallow water
bodies only (< 1.8 m).
Mats of shallow lakes (epipsammic and flake mats) and those of
deeper lakes (prostrate mats) were characterized by different dominant

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 202 A. TATON et al.

cyanobacterial morphotypes. In contrast to diatoms, no relationships

between the cyanobaterial morphotypes and conductivity were found.
Mats of shallow lakes (flake mats) were characterized by the presence
of Nostocales while the deeper lake mats where largely dominated by
Oscillatoriales (SABBE et al. 2004).
The matrices of the mats, and thus their structures, were built by the
filamentous taxa, mostly thin Oscillatoriales, whereas Chroococcales were
growing in small patches in this matrix.
Leptolyngbya was the most abundant morphotype in our samples. It
was frequently and abundantly found in prostrate mats and not found in
epilithic mats. This taxon probably included several species that were not
distinguished due to the complexity of the mat samples and the fact that
the trichomes of this genus are extremely thin. Phormidium priestleyi and
Pseudanabaena frigida were also frequently found in our samples; both
mostly in prostrate mats. Phormidium priestleyi was found in epipsammic
mats whereas Pseudanabaena frigida was found in epilithic and finely
lamellate mats. Pseudanabaena cf. amphigranulata and Pseudanabaena cf.
tremula were only found in prostrate mats, except one flake mat sample
that contained Pseudanabaena cf. tremula. Schizothrix lacustris and
Schizothrix cf. simplicior were characteristic of flake mats though they
were occasionally found in the other mat types, except the epilithic mats.
Schizothrix cf. tenuis was abundantly found in one epilithic mat sample.
The Nostocales were mostly found in flake mats. However, Nostoc sp.
was present throughout all the mat types. Petalonema cf. involvens and
Coleodesmium cf. scottianum were rarely found and limited to flake mats;
Dichothrix sp. and especially Calothrix cf. parietina were more often found,
though not in abundance. Among the Chroococcales, Chlorogloea sp. was
the most often found and the most abundant taxon. The distribution of
Chondrocystis cf. dermochroa is characteristic, since this taxon was quite
abundant in flakes and rarely observed in the other mat types. Asterocapsa
sp., Chamaesiphon subglobosus, Gloeocapsa cf. alpina, Gloeocapsa cf.
compacta, Gloeocapsa cf. sanguinea, Rhabdonema sp. and cf. Chamaesiphon
were rarely found. Members of the genera Aphanothece and Aphanocapsa
were mainly recorded in prostrate mats. However, Aphanothece cf. hyalina
was only found in lake mats. The identification of several unicellular taxa
outlined the importance of such microscopic studies compared to molecular
ecology studies, as the latter approach has failed to detect these organisms
in similar samples (TATON et al. 2006a).
Several species found in our samples of mats of the Larsemann Hills
lakes were commonly found in the periphyton and the mats of lakes in other
regions: Chondrocystis cf. dermochroa, Gloeocapsa cf. alpina, Chamaesiphon
subglobosus, Pseudanabaena frigida, Phormidium priestleyi, Phormidium
autumnale, Phormidium murrayi, Calothrix cf. parietina. However, none
seems to be restricted to the periphyton of lakes and ponds. At least 7 of

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 Cyanobacteria in mats of East Antarctic lakes 203

Fig. 13. Relative abundances (circle diameter) of each morphotype (x-axis) found
in the studied lakes (y-axis) (for the abbreviation see tables 1 and 2).

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 204 A. TATON et al.

them were even recorded from terrestrial environments. KOMÁREK (1999)

emphasized the importance of the ecological niche in the identification of
Antarctic cyanobacteria. A question that can be addressed is therefore: Do
morphologically similar cyanobacteria found in different ecological niches
belong to the same or different species?
It is obvious that morphology does not always reflect the physiological
and genetic diversity (Vincent 2000a). The success of cyanobacteria in
Antarctica is well recognized and attributed to their tolerance to desiccation,
freeze thaw cycles, intense solar radiation, UV… (e. g. HAWES et al. 1999,
TANG et al. 1997, VINCENT & QUESADA 1994). Different taxa should or
could be more or less tolerant to particular conditions. However, GORDON
et al. (2000) have demonstrated, by using oligonucleotide probes specific for
16S rRNA gene sequences of uncultured Antarctic cyanobacteria isolated
from the ice cover of Lake Bonney, that the lake ice microbial community
was dominated by organisms that originated elsewhere in the surrounding
region (e. g. terrestrial microbial mats). Therefore, some taxa (recorded
in the present or other studies) may have been found in a transitory
environment of their colonization process.
Another important topic concerns the biogeography of Antarctic
cyanobacteria. Most taxa were previously found outside Antarctica. Only
two species, Pseudanabaena frigida and Phormidium priestleyi, would be
endemic to Antarctica (KOMÁREK 1999, KOMÁREK & ANAGNOSTIDIS 2005).
As previously shown, morphological results contrasted with 16S rRNA
based studies of Antarctic microbial mats. Indeed, molecular data suggested
that most of cultivated and uncultivated cyanobacteria from these biotopes
were potentially endemic or novel (TATON et al. 2003, 2006a, 2006b).
Twelve taxa (Gloeocapsa cf. compacta, Asterocapsa sp., Aphanothece
cf. elabens, Rhabdonema sp., Aphanocapsa cf. holsatica, Aphanocapsa cf.
hyalina, cf. Jaaginema sp., Pseudanabaena cf. tremula, Schizothrix cf. tenuis,
Schizothrix cf. simplicior, Coleodesmium cf. scottianum and Petalonema
cf. involvens) were found for the first time in Antarctica. Beside these 12
newly recorded taxa for Antarctica and the 5 taxa determined at the genus
level, the others were found in at least one other Antarctic region. Potential
endemic species (Pseudanabaena frigida and Phormidium priestleyi) as
well as the (sub-)cosmopolitan Phormidium autumnale and Gloeocapsa cf.
sanguinea are particularly widely distributed over the continent. Molecular
ecology studies have also revealed a “wide” distribution of a number of
taxa over the Antarctic continent, particularly (sub-)cosmopolitan taxa.
These taxa probably more adapted to transportation and dissemination
should have been quite successful in spreading to new Antarctic habitats
in different regions following Holocene deglaciation and isostatic uplift
(TATON et al. 2006a).
Interestingly, 8 taxa (Gloeocapsa cf. compacta, Gloeocapsa cf. alpina,
Gloeocapsa cf. sanguinea, Asterocapsa sp., Chamaesiphon subglobosus,

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 Cyanobacteria in mats of East Antarctic lakes 205

Aphanocapsa cf. hyalina, Schizothrix lacustris, Schizothrix cf. simplicior),

also found outside Antarctica, were characteristic for or at least were
previously found in the subaerophytic communities or in the freshwater
environments of mountains where the temperatures are usually lower than
in lowlands, particularly during the winter. Furthermore, several species
are also known from Arctic regions, including Pseudanabaena cf. tremula
recently described by CASAMATTA et al. (2005). Molecular ecology studies
of benthic communities in high altitude lakes and ponds are currently
lacking. Such studies would permit to draw out more definitive conclusions
concerning the biogeography of psychrotolerant cyanobacteria.

Concluding remarks
This study has brought new knowledge concerning the cyanobacterial diver-
sity in microbial mats of East Antarctic Lakes. However, due to problems
encountered in the cyanobacterial identifications, very interesting questions
in relation to the ecology and biogeography of Antarctic cyanobacteria re-
main unanswered and need the use of molecular tools. Such a molecular
diversity study has been started for several samples from lakes and ponds
of the McMurdo Ice Shelf, Dry Valleys, Vestfold Hills and Larsemann Hills
(e. g. JUNGBLUT et al. 2005, TATON et al. 2003, 2006a). Data showed that the
genetic diversity, based on the 16S rRNA sequences, was higher than the
morphological diversity and brought more reliable answers concerning
ecology and biogeography of Antarctic cyanobacteria. Nevertheless, they
also outlined the usefulness of more traditional morphological approaches.

Acknowledgments

This study was funded by the European Union Biotechnology Program through the
MICROMAT project (grant BIO4-CT98-0040). ANNICK WILMOTTE is a research
associate of the National Fund for Scientific Research (Belgium). ARNAUD TATON
had a fellowship from the Funds for Research Formation in Industry and Agri-
culture (Belgium). We thank PATRICK MOTTE (Biological Sciences, University of
Liège) who kindly lent us a digital camera equipped microscope. We thank DOMINIC
HODGSON and PHILIPPA NOON (British Antarctic Survey), who collected the mat
material and organized the transport of samples. We wish to express our gratitude
to JIŘÍ KOMÁREK who helped us for the identification of small unicellular taxa.

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Manuscript received January 12, 2007, accepted September 15, 2007.

The authors’ addresses:

Dr. ARNAUD TATON

Dr. ANNICK WILMOTTE
Centre d’Ingénierie des Protéines
Institut de Chimie B6, Université de Liège
B-4000 Liège, Belgium
E-mail: awilmotte@ulg.ac.be

Dr. LUCIEN HOFFMANN

Department ‘Environment and Agro-biotechnologies’
Centre de Recherche Public-Gabriel Lippmann
L-4422 Belvaux, Grand-duchy of Luxembourg

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