Br. phycol. J.

11 : 387-405
1 December 1976

S I X N E W S P E C I E S OF T E R R E S T R I A L A L G A E
FROM SIGNY ISLAND, SOUTH ORKNEY
ISLANDS, ANTARCTICA
By PAUL A. BROADY*
British Antarctic Survey, Life Sciences Division, Madingley Road, Cambridge, CB3 0ET

Six species are described and compared with related algae. All were isolated in culture from
terrestrial habitats on Signy Island, South Orkney Islands, Antarctica. They are: Botrydiopsis
constricta sp. nov. (Mischococcales, Xanthophyceae), Heterothrix antarctica sp. nov. (Tribo-
nematales, Xanthophyceae), Sphaerocystis oleifera sp. nov. (Chlorococcales, Chlorophyceae),
Sphaerocystis signiensis sp. nov., Sphaerocystis bilobata sp. nov. and Fottea pyrenoidosa sp. nov.
(UIothricales, Chlorophyceae).
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The algae described in this p a p e r were f o u n d d u r i n g a survey o f terrestrial

algae conducted at the research station of the British Antarctic Survey on Signy
Island, South O r k n e y Islands (60 ° 43' S, 45 ° 38' W) between 1971 a n d 1974.
Signy Island lies in the m a r i t i m e Antarctic as defined by Holdgate (1964) a n d
has been described in some detail by Holdgate (1967).
The study of the terrestrial algae included an e x a m i n a t i o n , by culture a n d
direct observation, of a wide range of terrestrial sites. Lists a n d descriptions o f
the sites examined, the algae found, a n d their d i s t r i b u t i o n are given by Broady
(in press). The following descriptions of algae new to science result f r o m further
work o n cultures b r o u g h t back to the U n i t e d K i n g d o m a n d studied in the
D e p a r t m e n t of Botany, University o f Bristol.
Six new species are proposed. T w o species, Botrydiopsis eonstricta sp. nov.
and Heterothrix antarctica sp. nov., are members of the Xanthophyceae. The
remaining four, Sphaerocystis oleifera sp. nov., S. signiensis sp. nov., S. bilobata
sp. nov. a n d Fotteapyrenoidosa sp. nov. are all members of the Chlorophyceae.

MATERIALS AND METHODS

Samples of soil and vegetation were collected using aseptic techniques and were transported,
in sterile containers, to the laboratory at the research station on Signy Island. Sample sites
included bare mineral soil, carpets, turves and cushions of several species of moss, soil from
below some small stands of herbaceous vegetation, and biotic soils formed from mud, seal and
penguin faeces and moulted hair and feathers. The samples were examined within 24 h of
collection. Moist plate enrichment cultures (Lund, 1945) were set up and were examined
frequently over a period of 26 weeks. Algae were also cultured on mineral agar medium. The
innoculum for these cultures consisted of untreated sample material, material from the moist
plate enrichment cultures, homogenized sample material in sterile water, or dilutions of the
homogenized sample. The latter treatment was performed at 14,000 r.p.m, on an M.S.E.
homogenizer and the dilutions of the homogenates were used for making culture counts.
Bold's modified Bristol's medium (BBM) (Chantanachat & Bold, 1962) was used for all the
culture work. Oxoid agar No. 3 at a concentration of 2 ~ w/v was used for plate cultures.
Incubation was at room temperature (c. 18°C) in constant light supplied by 4 × 30 W daylight
fluorescent tubes (Cryselco Ltd.). Individual colonies appearing in culture were transferred to
BBM agar slopes in McCartney vials. A collection of unialgal cultures was maintained by
* Present address: Department of Botany, University of Bristol, Bristol BS8 1UG.
387
 388 P . A . BROADY

periodic transfer of the cultures onto fresh slopes. For transport to the United Kingdom the
cultures were kept deep frozen at --20~C. They remained at this temperature for 1 year. On
thawing they were innoculated onto fresh BBM 2 ~ agar slopes, with 15 ml of garden soil
extract added in each litre of medium. Cultures were also grown in aqueous medium. Frequent
microscopic examinations were made on material from these cultures using a Wild M20
microscope with magnification up to ~\ 1000. The presence of starch granules, pyrenoids, the
shapes of the chromatophores, and the number of nuclei were made more readily visible with
Lugol's iodine solution. Indian ink and methylene blue were used to investigate the presence
of mucilaginous secretions. Sudan 4 was used to stain for oil.

OBSERVATIONS AND DISCUSSION

BOTR Y D I O P S I S C O N S T R I C T A SP. N O V .
DIAGNOSIS
Cellulae maturae plerumque sphaericae, vel interdum late ellipsoideae, singulae, ad 42 t~m
diam., plurimae cellularum majorum c. 25 ~m diam. Paries cellulae taevis, tenuis, aliquando
partibus incrassatis lentiformibus. Cellulae magnae chromatophoris multis, plerumque
fusiformibus, sine pyrenoidibus, plurimis parietalibus, etsi saepe nonnullae vel in cellulam
procurrunt vel in interiore cellulae sunt. In cellulis juvenibus ad 10 t~m diam. chromatophora
pauca, parietalia, disciformia. Cellulae maturae multinucleatae, nucleis ad centrum cellulae.
Globuli oleosi saepe adsunt. Interdum, in culturis vetustis, cellulae singulae vagina stratosa
gelatinosa circumcinctae. Multiplicatio aplanosporis, zoosporis et divisione vegetativa.
Aplanosporae plerumque sphaericae, 3-5-6 t~m diam., sed interdum ellipsoideae ad fete sub-
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cylindricae, uno, duobus, vel, rarenter, tribus vel quattuor chromatophoris, uninucleatae.
Zoosporae 5-7x2-3/~m, nudae, inaequaliter biflagellatae, unaquaeque chromatophoro
singulo, stigma capienti, plerumque multae simul ruptura parietis cellulae liberatae. Divisio
vegetativa plerumque in culturis vetustis; cellulae sphaericae, plerumque majores quam
20 t~m diam., late ellipsoideae ad fete cylindricae fiunt, cellulae constringtmt et, formatione
parietis transversalis trans constrictionem, duae cellulae filiales aequales vel iaequales formantur.
Reproductio etiam velut pullulando formatione duarum cellularum filialium inaequalium.
Formae cellularum tempore divisionis vegetativae multae et irregulares.
Ex solo et muscis in "Signy Island, South Orkney Islands, Antarctica" culta, 1972 lecta.
Holotypus: Figurae 1 a-s and 2 a-q.
Adult cells usually spherical, or broadly ellipsoidal, single, up to 42 p.m diam., majority of
larger cells about 25/zm diam. Smooth, thin cell wall, occasionally with lens shaped thicken-
ings. Large cells with many chromatophores, usually spindle-shaped, lacking pyrenoids, mostly
parietal though often some either project into the cell or are entirely in the interior. Young cells
up to 10 tzm diam. with a small number of parietal, plate-like chromatophores. Adult cells
multinucleate with the nuclei in the centre of the cell. Oil globules often present. Occasionally,
i n old cultures, individual cells surrounded by a stratified gelatinous sheath. Multiplication by
aplanospores, zoospores and vegetative division. Aplanospores usually spherical, 3"5-6/~m
diam., occasionally ellipsoidal to almost cylindrical with one, two or rarely three or four
chromatophores and a single nucleus. Zoospores, 5-7 × 2-3 t~m, naked, unequally biflagellate,
with a single chromatophore containing a stigma; usually released in large numbers by the
rupture of the sporangium wall. Vegetative division most frequently seen in old cultures.
Spherical cells, usually greater than 20 p.m diam., becoming broadly ellipsoidal to almost
cylindrical, the cells constricting and a transverse wall forming across the constriction thus
cutting off two daughter cells of equal or unequal sizes. A process resembling budding also
observed giving rise to two unequal daughter cells. Cells in the process of vegetative division
displaying many irregular shapes.
Cultured from soil and moss, Signy Island, South Orkney Islands, Antarctica, 1972.
Holotype: Figs 1 a-s and 2 a-q. Material deposited at the Cambridge Collection of Algae
and Protozoa as Culture No. 806/4.

N o s t a r c h o r o t h e r g r a n u l e s are r e v e a l e d in t h e c h r o m a t o p h o r e s b y t r e a t m e n t
w i t h L u g o l ' s iodine. O i l g l o b u l e s a r e r e a d i l y i d e n t i f i e d u s i n g S u d a n 4 stain.
S o m e a d u l t cells b e c o m e p a c k e d w i t h oil.
Z o o s p o r e s are f o r m e d w h e n m a t e r i a l f r o m o l d a g a r c u l t u r e s is r e i n o c u l a t e d
i n t o fresh g r o w t h m e d i u m . T h e y are also f o r m e d in l i q u i d c u l t u r e s u p to 2 m o n t h s
old. I n t h e first stage o f z o o s p o r e p r o d u c t i o n a n o r a n g e - r e d s t i g m a a p p e a r s in
e a c h c h r o m a t o p h o r e (Fig. 1 l - n ) . T h i s o c c u r s in small cells, o n l y 8 / ~ m d i a m . ,
 New algae from Antarctica 389

i. .................................... j...:ii:i.iis ~
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FIG. I. Botrydiopsis constricta. (Scale 10 t~m except p-r.) a-d, young cells; e, ellipsoidal
cell showing multinucleate condition; f, g, large adult ceils; h, adult cell with chromato-
phores radially arranged and closely packed; i, j, gelatinous sheaths round cells; k, a
cell wall thickening; l-n, zoosporangia at various stages of development, n, showing
transverse section and surface views; o, ruptured zoosporangium; p, zoospore swim-
ming; q, zoospore settled and putting out pseudopodia; r, zoospore after withdrawing
flagella; s, autosporangium.
 390 P.A. BROADY

g
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FIG. 2. Botrydiopsis constricta. (Scale 10/zm.) a, early vegetative division; b-c, late stage
in division; d, prior to cell separation; e, commencement of "budding" division: f, later
stage; g, constriction and formation of a small daughter cell; h-q, camera lucida draw-
ings of cells from an actively dividing culture, showing cells where wall thickenings were
present.
 New algae from Antarctica 391
with discoid chromatophores, in large cells with spindle-shaped chromatophores
and even in cells undergoing vegetative division. The chromatophores often take
up positions with their long axis radially disposed and with the stigmata lying
close to the cell wall (Fig. 1 n). The zoospores are then formed in large numbers
by partition of the protoplast. A sporangium only 10 pm wide contains over
twenty spores and one 25 #m wide contains over a hundred. They are released
by the irregular rupture of the sporangium wall which remains visible amongst
the living cells. The rupture is usually only limited in extent (Fig. 1 o). The active
zoospores swim with a corkscrew motion for several minutes before settling.
They can then be seen to lack a rigid cell wall when pseudopodia develop
(Fig. l q). The spores soon round off, withdraw their flagella, and lose the two
contractile vacuoles (Fig. 1 r). The flagella are inserted slightly laterally, one
long easily viewed flagellum stretches out to the front of the swimming spore
and a short flagellum, viewed with difficulty, trails towards the rear (Fig. 1 p).
The stigma usually lies in an anterior position.
Aplanospores are formed mostly in young cultures before the onset of
vegetative division. Recently released spores may remain in loose aggregates
for a short while after release from the ruptured sporangium. Aplanospores with
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a stigma have been seen aggregated around a ruptured sporangium wall. These
could have been released in this condition or could have been zoospores which
had rapidly lost their flagella.
Vegetative division (Fig. 2) occurs mostly in old cultures, especially on agar.
A selection of camera lucida drawings of cells exhibiting various stages of
division are shown in Fig. 2 h-q. Many cells also remain spherical. Cells under-
going vegetative division are usually greater than 20 pm diam. Division takes
place in two ways. Firstly, spherical cells may become broadly ellipsoidal to
almost cylindrical. A constriction appears, usually in a median position, and the
cell wall in this region often thickens and becomes faintly lamellate (Fig. 2 a).
Thickenings are not always present (Fig. 2 i, m and q). The constriction becomes
deeper (Fig. 2 b) and the cell wall thickenings closer together until the original
protoplast is divided into two (Fig. 2 h, o). A thickish bridge of cell wall material
may be present between the two daughter cells before they separate (Fig. 2 d).
The daughter cells are sometimes of equal diameter and sometimes differ
markedly (Fig. 2 h). In the second type of division, which usually leads to two
cells of unequal size, the mother cell forms a bud which developes to form a
daughter cell. Bud initiation commences with the formation of a thickening of
one portion of the cell wall (Fig. 2 e). The mother cell becomes pyriform
(Fig. 2 f) and the daughter bud then swells and becomes separated from the
mother cell by a deepening constriction (Fig. 2 g). The cell wall is usually
thickened at the point of constriction. Eventually cell wall material bridges the
constriction and separates off the two protoplasts. These then separate as two
daughter cells.
This alga was frequently found in mineral soils, in the wetter moss carpet
communities (Drepanocladus uncinatus (Hedw.) Warnst. and Calliergon
sarmentosum (Wahlenb.) Kindb. and in soil beneath the grass Deschampsia
antarctica Desv., although it was never the dominant alga.
This alga is clearly a member of the Xanthophyceae since it has yellow-green
chromatophores, starch is absent and the swarmer is unequally biflagellate.
 392 P.A. BROADY

Bourrelly (1968) describes three genera of free-living, non-motile, spherical

algae in this class, which also possess several parietal chromatophores lacking
pyrenoids. These are Pleurochloris Pascher, Chloridella Pascher and Botrydiopsis
Borzi. The first two genera have adult cells of a small size with only a few
chromatophores and Chloridella does not produce zoospores. The present alga
can be assigned to the genus Botrydiopsis because of the large adult cells with
numerous chromatophores which develop from much smaller cells with a small
number of chromatophores, the latter derived from zoospores or aplanospores
(Borzi, 1895).
Six species of Botrydiopsis have been reliably described. All of these reproduce
by the production of heterokont, biflagellate, naked zoospores, or aplanospores,
as does B. constricta. However, no mention is made of a vegetative division as in
the present alga. This is the major characteristic separating B. constricta from
other described species. The zoospores of B. constricta are also characteristic
(Table I), no other species has spores with a stigma and only one chromatophore.

TABLE I. The number of chromatophores and the presence or absence of a stigma

in zoospores of species of Botrydiopsis
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Number of Presence of
chromatophores a stigma (+)

B. arhiza 1-3 --
B. intercedens 2 4-
B. anglica 2 +
B. alpina 1-2 +
B. pyrenaica 2 --
B. antarctica 1-2 +
B. constricta 1 +

The vegetative cells of B. alpina Vischer, B. pyrenaica Viala and B. antarctica

Kol do not vary from spherical, unlike the present alga. B. antarctica is similar
to B. constricta in that the cells can form a thick, lamellar, gelatinous coat in
culture.
Other species possess vegetative features which are similar to those displayed
by B. constricta. B. arhiza Borzi has mostly spherical adult cells but some may
be ellipsoidal to cylindrical. Deason & Bold (1960) in an examination of a strain
of B. arhiza, isolated from soil, illustrate an almost cylindrical cell which is
slightly constricted centrally (their Fig. 123) in a similar manner to early stages
of vegetative division in B. constricta. They do not describe any extension of this
cell shape into a vegetative division despite examining the alga in culture on
Bristol's agar. The B. arhiza studied by Chadefaud (1943) produced pyriform
cells on occasions, similar to B. constricta at an early stage of vegetative division.
Again no mention is made of this process occurring. B. anglica (James) Fritsch
et John is illustrated in James (1935) as having occasional cells with a lamellate,
lens shaped, cell wall thickening as present in some cells ofB. constricta (Fig. 1 k).
However the adult cells of B. anglica differ in that they have a thick stratified
cell wall.
 New algae from Antarctica 393

H E T E R O T H R I X ANTARCTICA SP. NOV.

DIAGNOSIS
Filamenta flexuosa, multicellularia, in culturis juvenibus sine constrictionibus ad septa.
Unaquaeque cellula chromatophoro singulo, luteo-viridi, parietali, disciformi, dimidium
parietis interioris cellulae tegens. In ceUula dividenti duo chromatophora facta sunt antequam
septum factum est. In filamentis juvenis chromatophorum uniuscuiusque cellulae tam longum
quam cellula, sed in unaquaeque cellula longiore in filamentis vetustis chromatophorum ad
centrum. Globuli oleosi saepe adsunt, praecipue in culturis vetustis. Filamenta 3-4-5 t~m lata,
cellulae in culturis vetustis 3"5-9 t~m, in culturis juvenibus ad 18 t~m, Iongae. In filamentis
vetustis septa interdum concavo-convexa. Akineta in eulturis vetustis prompte facta, plerumque
sphaerica, 3'5-10 tLm, rarenter ad 15/~m, diam., unumquidque pariete saepe incrassato spinis
brevibus rectum.
Ex cultura ex muscis terrestribus in habitatione humida, acida in "Signy Island, South
Orkney Islands, Antarctica" segregata, 1972 lecta.
Holotypus; Figura 3 a-m.
Long, multicellular flexuous filaments, in young cultures without constrictions at the cross-
walls. Each cell with a single, yellow-green, parietal, plate-like chromatophore covering about
half the inner cell wall. Dividing cells with two chromatophores before formation of the
transverse wall. Chromatophores equal to cell length in young filaments but situated centrally
in cells of older filaments. Oil globules often present particularly in older cultures. Filament
width 3-4.5 tzm, cell length 3"5-9 vm in young, and up to 18 t~m in old cultures. In old filaments
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the transverse walls sometimes concavo--convex. Akinetes readily formed in old cultures,
usually spherical, 3"5-10 t~m, rarely 15 tzm diam., often with a thickened wall covered with
short spines.
Isolated in culture from terrestrial mosses in a wet, acid habitat on Signy Island, South
Orkney Islands, Antarctica, 1972).
Holotype: Fig. 3 a m . Material deposited at the Cambridge Collection of Algae and
Protozoa as Culture No. 836/6.

Fragmentation occurs in b o t h y o u n g and old cultures but is far m o r e frequent

in the latter where it may be extensive and followed by the formation o f m a n y
akinetes. At the first stage o f fragmentation (Fig. 3 c) constrictions appear at
the cross walls and short lengths o f filament break away. These m a y then split
into individual vegetative cells (Fig. 3 d, e). Occasionally short lengths o f
filament m a y fragment into single cells without first splitting into shorter but
multicellular lengths. Such filaments have constrictions at every transverse wall
prior to their disintegration (Fig. 3 k). The single vegetative cells become
spherical (Fig. 3 f), sometimes ellipsoidal or subspherical. The cell wall then
becomes sculptured with short warty protuberances (Fig. 3 g-j) and often
increases to about 1 # m in thickness. These akinetes are usually packed with oil
droplets and contain a single, parietal, somewhat lobed chromatophore. Some-
times a large oil globule almost fills the resting cell (Fig. 3 i). On innoculation o f
material containing akinetes into fresh medium they germinate directly into the
typical filaments and no akinetes are found in the y o u n g culture. Rarely a
vegetative cell still attached to a short filament may f o r m the sculptured cell wall
(Fig. 3 1).
Staining o f the cells with Lugol's iodine does not reveal the presence of starch.
However, golden-brown staining granules are scattered t h r o u g h o u t the chro-
matophore (Fig. 3 m), these are possibly chrysolaminarin. The single, central
nucleus is also made more apparent.
This alga was infrequently isolated from cultures o f moss and the underlying
peat from wet, acid carpets o f Drepanocladus uncinatus and Calliergon
sarmentosum.
 394 P.A. BROADY
a

d
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I •

FrG. 3. tteterothrix antarctica. (Scale 10 ~m except in.) a, young filament; b, old fila-
ment; c, fragmentation commencing; d, e, later stage of fragmentation; f, rounded
vegetative cell, early akinete formation; g, h, fully formed akinetes; i, large oil globule
in akinete; j, surface view of akinete; k, short filament in which all cells starting to
fragment; 1, akinete formation whilst still attached; m, iodine stained cell showing
golden-brown granules and nucleus.
The yellow-green chromatophores and the absence o f starch place this alga
in the Xanthophyceae. It has the characters o f Heterothrix Pascher in which the
cell walls, in the vegetative state, cannot be recognized as being composed o f
two half cells as is the case with the related genera Tribonema Derbes and Solier
and Bumilleria Borzi (Pascher, 1932).
Sixteen species have been described. 11. antarctica is c o m p a r e d with similar
species in Table II. The nine species not included in the table are unlike H.
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TABLE II. A comparison of Heterothrix antarctica sp. nov. with similar species

Length of filaments
L: Long Constriction o f
F: Short, Zoospore Akinete y o u n g filaments
Filament Cell length easily production production Number of at cross walls
width (t~m) (tzm) fragmented (+) (+) chromatophores (+) Habitat

H. antarctica 3.0-4-5 3-5-9.0 L -- _ 1 -- Terrestrial

sp. nov. ( - - 18'0) mosses
H. monoehloron 2"0-3'0 -4"5 L + -- 1 ÷ Freshwater
Ettl
H. monochloron 2'3-3"7 4'6--11"1 L -- -- 1 + Soil
var. terrestre
Ettl et K~cha
H. stichocoeeoides 2'0-3'0 --8'0 F + + 1 + Soil
Pascher
H. hormidioides 4"0 4-0-10-0 L -- -- 1-2 + Soil
Vischer
H. exilis 3"5-4"5 6"0-8"0 F -- ÷ 2-t- -k Soil
Pascher
H. ulotrichoides 9"0-12-0 --16"0 F ÷ + 2 + Soil
Pascher
H. Bristoliana 4"0-6"0 10"0-15"0 F + + 4-12 -k Soil
Pascher
H. fluviatilis 7"0 7-0-15'0 L + -- 1 -k Soil
Gayral et
Mazancourt
 396 P.A. BROADY

antarctica in that they do n o t p r o d u c e akinetes, they have two or m o r e c h r o m a t o -

p h o r e s in each cell, a n d t h e y usually p r o d u c e z o o s p o r e s o r occasionally a p l a n o -
spores. O f the species in the table, H. monochloron var. terrestre Ettl et K a c h a
a n d H. hormidioides Vischer are m o s t similar to H. antarctica. The r e m a i n d e r
p r o d u c e zoospores, some have m o r e t h a n one c h r o m a t o p h o r e , a n d f o u r species
f o r m short, easily f r a g m e n t e d filaments. However, n e i t h e r o f the two similar
species p r o d u c e akinetes, a n d filaments o f b o t h have constrictions at the
transverse cell walls, unlike H. antarctica in y o u n g cultures. Because o f these
differences H. antarctica is p r o p o s e d as a new species.

SPHAEROCYSTIS S I G N I E N S I S SP. N O V .
DIAGNOSIS
Cellulae singulae, sphaericae vel rarenter late ellipsoideae, ad 15/~m diam., in coloniis molle
mucosis, multicellularibus, irregularibus, atrovirentibus, ubique binae, quaternae et, rarenter,
octonae aggregatae vel irregulariter dispositae. Circum cellulas et greges cellularum interdum
structura inconspicua muci. Cellula matura pariete tenui, laevi, chromatophoro singulo, saepe
lobato, parietali, viridi, pyrenoide basali evidenter visibili, et nucleo singulo. Globuli oleosi
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Fie. 4. Sphaerocystis signiensis. (Scale I0/~m except a and k.) a, cells in colony; b-e,
young cells recently released from sporangium; f, adult cell; g, large vacuoles in adult cell;
h, iodine stained cell with starch granules and nucleus; i, j, cells at an early stage of
division; k, faint structuring of mucilage.
 New algae from Antarctica 397

saepe adsunt. Autosporangia 2, 4 vel interdum 8 autosporis sphaericis vel late ellipsoideis,
4"5-7 tzm diam., ruptura irregulari parietis sporangii, cujus reliquiae saepe visibiles, vel
gelatinisatione ejus liberatis.
Ex muscis terrestribus in "Signy Island, South Orkney Islands, Antarctica" culta, 1972 lecta.
Holotypus: Figura 4 a-k.
Spherical, rarely broadly ellipsoidal, unicells, up to 15/~m diam. distributed throughout
softly mucilaginous, multicellular, irregular, dark green colonies in two, four and rarely eight
cell groups, or irregularly arranged. Occasionally with a faint structuring of the mucilage
around cells and cell groups. Adult cells with a thin, smooth cell wall, and a single, often lobed,
parietal, green chromatophore with a readily visible basal pyrenoid, and a single nucleus. Oil
globules often present. Autosporangia release two, four rarely eight spores by the irregular
rupture of the wall, fragments of which being often visible, or gelatinizing. Spores spherical or
broadly ellipsoidal, 4"5-7 t~m diam.
Isolated in culture from terrestrial mosses, Signy Island, South Orkney Islands, Antarctica, 1972.
Holotype: Fig. 4 a-k. Material deposited at the Cambridge Collection of Algae and Protozoa
as Culture No. 176/3.

S P H A E R O C Y S T I S OLEIFERA SP. N O V .
DIAGNOSIS
Cellulae singulae, late ellipsoideae vel subsphaericae, ad 11 p.m diam., in coloniis molle
mucosis, multicelularibus, irregularibus, subviridibus, ubique binae, quaternae, octonae vel
senae denae aggregatae; cellulae magnae maturae interdum irregulariter dispositae. In muco
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oo0 ®
o( o
© d

........
• .. ................... ............................ ~.

!0

FIG. 5. Sphaerocystis oleifera. (Scale 10 ~m except a and j.) a, cells in colony; b-e, young
cells recently released from sporangium showing range of chromatophore shape; f, g,
adult cells; h, i, iodine stained cells with nuclei and starch grains;j, faint structuring of
mucilage.
G
 398 P . A . BROADY

circa greges cellularum interdum strata concentrica vixvisililes. Cellula matura pariete tenui,
laevi, in culturis vetustioribus ad 1"5 tLm parum et aeque spissescenti, nucleo singulo et
chromatophoro singulo, parietali, late lobato, viridi, pyrenoide vixvisibili. Autosporangia 4,
8 vel, rariore, 2 vel 16, autosporis, ruptura irregulari parietis sporangii liberatis. Reliquiae
parietis sporangii aliquando visibiles remanentes, aliquando deliquescentes. Sporae juvenes
anguste ellipsoideae ad fere cylindricae, 7-9 x 3-4 #m, unaquaeque chromatophorum parietale,
variabile lobatum continens. Globuli oleosi et in cellulas maturas et in autosporas plerumque
adsunt.
Ex culturis muscarum et soli in "Signy Island, South Orkney Islands, Antarctica" segregata,
1972 lecta.
Holotypus: Figura 5 a-j.
Broadly ellipsoidal to subspherical unicells, up to 11/~m diam., distributed throughout softly
mucilaginous, multicellular, irregular, light green colonies in groups of two, four, eight and
sixteen cells. Large adult cells occasionally irregularly distributed. Occasional faint, concentric
stratifications present in the mucilage around cell groups. Adult cells with a thin, smooth cell
wall, becoming slightly uniformly thicker, up to 1-5 tLm, in old cultures, containing a single
nucleus and a single green, parietal, broadly lobed chromatophore with a faint pyrenoid.
Autospores, four, eight and more rarely two or sixteen, released by irregular rupture of the
sporangium. Fragments of the sporangium wall may remain visible or become gelatinous.
Young spores narrowly ellipsoidal to almost cylindrical, 7-9 x 3-4/~m, with a variably lobed,
parietal chromatophore. Oil globules usually present in both adult cells and autospores.
Isolated in cultures of moss and soil, from Signy Island, South Orkney Islands, Antarctica,
1972.
Holotype: Fig. 5 a-j. Material deposited at the Cambridge Collection of Algae and Protozoa
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as Culture No. 176/2.

S P H A E R O C Y S T I S B I L O B A TA SP. N O V .
DIAGNOSIS
Cellulae sphaericae vel parum ellipsoideae, ad 8-5/zm sed plerumque 6 t~m diam., ubique in
coloniis multicellularibus, irregularibus, mucosis, atrovirentibus, quaternae vel octonae
aggregatae. Limites muci inter greges cellularum saepe visibiles quod bacteria et particulae
sordis in superficiebus suis adsunt. Cellula quaeque chromatophoro singulo, viridi, saepe
bilobato, extenso, parietali, partem maximam parietis interioris cellulae tegenti et pyrenoidem
basalem indistinctam inter lobos continenti. Nucleus singulus. Globuli oleosi saepe adsunt.
Autosporae 4 vel 8, 3 x 3'5 t~m, plerumque gelatinisatione parietis sporangii, vel rarenter
ruptura irregulari ejus, quum reliquiae circum unumquemque gregem sporarum visibiles sunt,
liberatae.
Ex culturis muscorum et soli turfosi ex stragulo humido, acido (pH 5'0) Calliergonis
sarmentosi (Wahlenb.) Kindb. segregata, in "Signy Island, South Orkney Islands, Antarctica"
1972 lecta.
Holotypus: Figura 6 g-l.
Spherical and slightly ellipsoidal unicells, up to 8-5 ~m but usually 6/~m diam., distributed
throughout multicellular, irregular, mucilaginous, dark green colonies, in four and eight cell
groups. The boundaries of the mucilage between each cell group often visible because of the
presence of bacteria and debris on their surfaces. Each cell with a single, green, often bi-lobed,
extensive, parietal chromatophore covering most of the inner cell wall and containing an
indistinct basal pyrenoid between the lobes. Oil globules often present and cells uninucleate.
Four and eight slightly eltipsoidal autospores, 3 x 3-5 t~m, usually released by gelatinization of
the sporangium wall, or by its irregular rupture, with fragments of wall remaining visible
around each spore group.
Isolated in cultures of moss and peat from a wet, acid (pH 5'0) carpet of Calliergon
sarmentosum (Wahlenb.) Kindb., Signy Island, South Orkney Islands, Antarctica, 1972.
Holotype: Fig. 6 g-l. Material deposited at the Cambridge Collection of Algae and Protozoa
as Culture No. 176/1.

T h e p a l m e l l o i d a l g a e o f t h e C h l o r o c o c c a l e s are a n o t o r i o u s l y difficult g r o u p
t a x o n o m i c a l l y . F o t t (1974) has a t t e m p t e d t o r e m e d y this s i t u a t i o n f o r g r e e n ,
u s u a l l y s p h e r i c a l o r e l l i p s o i d a l unicells in m u c i l a g i n o u s c o l o n i e s w h i c h u s u a l l y
reproduce by the formation of autospores and rarely by zoospores. Such forms
he has g r o u p e d in t h e n e w f a m i l y , P a l m o g l o e a c e a e . H e p r e s e n t s a k e y t o t h e
i d e n t i f i c a t i o n o f t h e g e n e r a a n d d e s c r i p t i o n s o f t h e species o f several o f these.
 New algae from Antarctica 399

The present algae all reproduce solely by autospores and are permanently
contained within mucilaginous colonies, they are clearly members of this family.
Using Fott's scheme four genera have characters close to the present algae;
Palmogloea Kuetzing, Coenochloris Korschikoff, Coenocystis Korschikoff and
Sphaerocystis Chodat. All these contain species with a single, parietal, usually
cup-shaped chromatophore, often containing a single pyrenoid. Characters of
the three new palmelloid algae are compared with the above four genera in
Table III.
The new species are excluded from Palmogloea because of their lack of clear,
concentric lamellations in the mucilage and by the presence of sporangium wall
remains (Fott & Nov~ikowl, 1971). In Coenochloris the autosporangium char-
acteristically splits into one or two portions which remain visible in the mucilage.
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FIG. 6. Sphaerocystis bilobata. (Scale 10 ~m except a.) a, cells in colony; b, bilobed

chromatophore in adult cell; c, surface view of cell showing the slight gap between the
chromatophore lobes; d, rare tri-lobed chromatophore; e, oil globules in cell; f, iodine
stained cell; g, h, dividing cells; i, auto-spores; j, autosporangium; k, 1, release of
autospores by k, gelatinization of the sporangium wall and 1, rupture.
 4~ P.A. BROADY

In all three new species more than two portions are often produced. The
a u t o s p o r a n g i u m wall o f Coenocystis usually gelatinizes and the adult cells are
never spherical. Only in Sphaerocystis bilobata sp. nov. do the remains o f the
sporangium wall usually gelatinize, but in this species the cells are usually
spherical which excludes it f r o m Coenocystis. Only in Sphaerocystis are similar
characters f o u n d as regards the shape o f the cells, the structuring of the mucilage
and the fate o f the sporangium wall. The three new species are thus assigned to
this genus.

TABLEIII. A comparison of the three new species of Sphaerocystis with the characters of
Sphaerocystis, and three similar genera as given in Fott (1974)

Remains of
Shape of Shape of Mucilage sporangium
adult cells colonies structure Reproduction wall

Palmogloea Subspherical, Irregular + Autospores --

ellipsoidal or 2, 4 and 8
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cylindrical
Coenochloris Spherical Spherical -- Autospores q-
ellipsoidal ellipsoidal 4, 8 rarely One or two
16 portions
Coenocystis Ellipsoidal, Spherical or -- Autospores --
never oval 4, 8 rarely
spherical 16
Sphaerocystis Spherical or Spherical 4- Autospores +
ellipsoidal ellipsoidal 2, 4, 8, 16 But may
or irregular rarely gelatinize
zoospores
S. oleifera Usually Irregular 4- 2, 4 rarely 8 q-
sp. nov. spherical autospores But may.
gelatinize
S. signiensis Subspherical Irregular 4- 2, 4, 8, 16 +
sp. nov. ellipsoidal autospores But may
gelatinize
S. bilobata Usually Irregular -- 4 and 8 --
sp. nov. spherical autospores May remain
visible

A comparison with the four previously k n o w n species, as described in F o t t

(1974) is given in Table IV. The original four species are distinguished by
differences in cell shape, habitat, the n u m b e r o f pyrenoids, and the presence or
absence o f zoospores. Each o f the three new species differs from each o f the
previously described species by several o f the characters in the table. In addition
the new species are also distinct, one f r o m another.
Only one o f the previously described species, S. bavarica, is f o u n d in a
terrestrial habitat, the remainder are planktonic and usually f o r m small colonies
o f few cells. The bilobed c h r o m a t o p h o r e o f S. bilobata is particularly distinctive
for that species, as are the narrowly ellipsoidal, almost cylindrical autospores o f
S. oleifera (Fig. 5 b ~ ) . S. signiensis is similar to S. schroeteri and S. bavarica.
It can be distinguished from the former by the shape o f the chromatophore and
b y the production o f two autospores. S. bavarica seldom has spherical cells and
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TABLE I V . A c o m p a r i s o n of the species of Sphaerocystis Chodat

Remains of
Colony shape Mucilage Size and shape Size and shape Chromatophore sporangium
and no. o f cells structure of adult cells of autospores shape Pyrenoid Reproduction wall Habitat

S. schroeteri 4-8-many cells Occasional Spherical Spherical Cup-shaped 1 4 , 8 o r 16 ± Planktonic

Chodat spherical or statifications 6-10 btm 4-5/zm with narrow basal autospores
broadly opening
ellipsoidal
S. planctonica 4-32 cells -- Spherical Broadly Cup-shaped 1 4, 8 seldom 16 Planktonic
(Korsch) spherical 9-12-5/~m ellipsoidal large basal autospores
Bourrelly also zoospores
S. polycocca 8-16 cells - - Spherical to Broadly Parietal 2-4 4, 8 autospores Planktonic
Korsch spherical or broadly ellipsoidal thick
broadly ellipsoidal
ellipsoidal 10-20/~m
S. bavarica Many cells Occasional Broadly Broadly Parietal 1 2,40r 8 q- Transition
(Skuja) irregular (Fort, 1974 cllipsoidal cllipsoidal occasionally autospores (Fott, 1974 moor
Bourrelly Fig. 5 a) seldom seen Figs 5 b-d)
spherical
6-15 ~ m
S. signiensis Many cells Occasional Usually Spherical or Cup-shaped 1 2, 4 occasionally + Terrestrial
sp. nov. irregular spherical broadly often deeply readily 8 autospores May mosses
7-15 ~ m cllipsoidal lobed visible gelatinize
4.5-7"0 p~m
S. oleifera Many ceils Occasional Subsphcrical Narrowly Parietal 1 2, 4, 8 or 16 + Terrestrial
sp. nov. irregular broadly cllipsoidal to broadly faint autospores May mosses and
ellipsoidal cylindrical lobed usually 4 or 8 gelatinize mineral soil
7-11/~m
S. bilobata Many cells - - Usually Broadly Parietal 1 4or8 Terrestrial
sp. nov. irregular spherical ellipsoidal usually faint autospores May remain moss in a
6-8"5/~m 3 × 3"5 I*m bilobed visible wet habitat
 402 P.A. BROADY

the pyrenoid is seldom seen, characters contrasting with the usually spherical
cells and the readily visible pyrenoid o f S. signiensis.
Although only one c h r o m a t o p h o r e was present in the normal vegetative cells
o f S. signiensis and S. bilobata, in early stages o f spore formation two, four or
eight parietal chromatophores were visible (Fig. 4 i, j, Fig. 6 h). In all three
species the presence or absence o f stratifications in the mucilage and remains o f
the sporangium wall were variable characters. Variations occurred with the age
and state of hydration o f the cultures. In old cultures there was a tendency for
the stratifications to be present and for the remains o f the sporangium walls to
be visible. Also in cultures which had dried out slightly the stratifications were
often more readily visible.
S. bilobata was only infrequently found. It was present in only one of the
seventy-two sites examined, namely the Signy Island terrestrial reference site 2
(Tilbrook, 1973) in an area consisting largely o f wet, acid carpets (pH 5.0) o f
the moss Calliergon sarmentosum. It was often recovered in cultures o f material
f r o m this one site and was a significant m e m b e r o f the algal flora. In contrast
the other two new species were f o u n d in a greater n u m b e r and a wider range o f
sites. S. signiensis was frequently isolated in cultures of moss and the underlying
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peat, particularly from; Polytrichum alpestre Hoppe, Chorisodontium aciphyllum

(Hook. f. et Wils.) Booth., Andreaea spp. and Calliergon sarmentosum. S. oleifera
is t h o u g h t to be a major constituent o f the algal flora o f Signy Island terrestrial
reference site 1 (Tilbrook, 1973) which consists predominantly of Polytrichum
alpestre and Chorisodontium aciphyllum. It was also f o u n d amongst Andreaea
spp. and Drepanocladus uncinatus as well as in bare, mineral soils.

FOTTEA P YRENOIDOSA SP. NOV.

DIAGNOSIS
Coloniae molle mucosae, multicellulares, irregulares. Cellulae cylindricae, plerumque parum
curvatae, rarenter rectae, apicibus late rotundatis, 6-18 t~m longae, 6 ~m latae, in muco
homogeneo irregulariter dispositae, aut singulae aut, in culturis dividentibus, duae, rarenter
tres, contingentes. Chromatophorum singulum, parietale, viride, disciforme, pyrenoidem
singulam continens, plus quam dimidium parietis interioris cellulae circumdans sed margines
ejus numquam conveniens. Nucleus singulus centralis. Globuli oleosi saepe adsunt. Divisio
cellularum vegetativarum in duas cellulas transversa.
Ex solo minerali (pH 5-2) in "Signy Island, South Orkney Islands, Antarctica" culta,
1972 lecta.
Holotypus: Figura 7 a-j.
Softly mucilaginous, multieellular, irregular colonies. Cells cylindrical, usually slightly
curved, rarely straight, with broadly rounded ends, length 9-I 8 t~m, width 6 ~m, irregularly
arranged throughout the homogeneous mucilage as single ceils or as two, rarely three, touching
cells in actively dividing cultures. Chromatophore single, parietal, green, plate-like, containing
a single pyrenoid, encircling over half the inner cell wall but never forming a complete inner
sleeve. Nucleus single central. Oil globules often present. Transverse division of vegetative
cells into two daughter cells.
In a mineral soil (pH 5"2), Signy Island, South Orkney Islands, Antarctica, 1972.
Holotype: Fig. 7 a-j. Material deposited at the Cambridge Collection of Algae and Protozoa
as Culture No. 326/I.
On agar the colonies are small, dark green and hemispherical but in liquid
culture small, irregular masses are formed (Fig. 7 a).
The single pyrenoid usually lies against the eoncave wall and the chromato-
phore doe s not cover the opposite convex wall. W h e n the cell is viewed looking
d o w n onto the pyrenoid, t h r o u g h the cell, it appears straight (Fig. 7 h). The
 New algae from Antarctica 403

chromatophore does not cover the end walls. Staining with Lugol's iodine
reveals starch grains encircling the pyrenoid and scattered throughout the
chromatophore, the single central nucleus is also made clearer (Fig. 7 i).
Transverse vegetative cell division is the only means of reproduction, spore
formation and resistant stages are absent. At an early stage in cell division the
chromatophore has two pyrenoids (Fig. 7 e), it then divides into two, each half
having a single pyrenoid (Fig. 7 f). The cross wall is then laid down and the cell
becomes more and more constricted at its midpoint until the two daughter cells

'": ~ %". :: ~0 o,::b ............~"r',..-,O "

:."d~ ._zzb % ..: ......... d::,: .......... c:~3 c.., c:b 0 .................

........................
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II j

FIG. 7. Fottea pyrenoidosa. (Scale 10 p.m except a.) a, cells in irregular colonies from
Liquid culture; b, young cell; c, spherical cross-section of cells; d, large cell; e-g,
vegetative division; h, view of cell looking down onto pyrenoid, cell appears straight;
i, iodine stained cell with starch and nucleus; j, rare short filament from an actively
dividing culture.
are separated off(Fig. 7 g). The ceils quickly part and there is usually no filament
formation. In actively dividing cultures two cells are often seen temporarily
touching and rarely a chain of three cells (Fig. 7 j) may be formed; many of the
cells also have two chromatophores. In stationary cultures most cells are single
and have just one chromatophore.
 4~ P.A. BROADY

This alga was only found in one of the seventy-two sites examined; fine,
mineral debris (pH 5-2) from a frost sorted stone polygon in an area of acid
quartz-mica-schist rocks. It was seen only in culture and never in examination
of field material.
The absence of contractile vacuoles, stigma and flagella in the vegetative state
and reproduction by the division of the mother cell into two parts, each incor-
porating the original mother cell wall, places this alga in the order Ulothricales
(Chlorophyceae).
The genus Fottea Hind~ik is thoroughly compared with similar genera by
Hind~ik (1968). The formation of gelatinous colonies in which cell distribution
is solitary and haphazard, as in the present alga, is one of the most distinctive
features of this genus. Elakatothrix Wille is close but here the dividing septum
in cell division becomes oblique and the daughter cells are spindle-shaped and
not cylindrical as in Fottea. The single, parietal chromatophore, the single
nucleus, the simple transverse division in one plane only and the absence of
zoospores, sexual and resistant stages are all additional characters which place
the present alga in Fottea.
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TABLEV. A comparison of Fotteapyrenoidosa sp. nov. with Fottea eylindrica HindS.k

Colony Cell size

shape Chromatophore Pyrenoid Cell shape (~m) Nucleus Habitat

F. cylindrica M a n y cells I -- Cylindrical 21-32 x 1 Soil, 2100 m

irregular parietal usually 8-12 altitude,
straight Czechoslovakia

F. pyrenoidosa M a n y cells I + Cylindrical 9-18 x 6 I Bare mineral soil,

irregular parietal usually 70 m,
curved Antarctica

F. pyrenoidosa is particularly unlike F. cylindrica, the only other known

species, in the presence of a pyrenoid. The two species are compared in Table V.
Although usually straight, the cells of F. cylindrica are sometimes slightly curved
similar to the usual form of F. pyrenoidosa.
Although the presence or absence of a pyrenoid is sometimes regarded as a
major character in delimiting genera (Starr, 1955), e.g. Dictyochloris Vischer and
Spongiochloris Starr of the Chlorococcales, in the present case the similarities of
F. cylindrica and F. pyrenoidosa are regarded as too great for the new species
to be placed in a separate genus. The closely related genus Elakatothrix also
includes species with and without pyrenoids (Hind~ik, 1962).

ACKNOWLEDGEMENTS
I wish to thank the British Antarctic Survey and its Director, Dr R. M. Laws, for providing
the opportunity to perform the field work on Signy Island. Professor D. C. Smith very kindly
provided facilities in the Department of Botany, University of Bristol, where subsequent work
was conducted. Dr F. E. Round, University of Bristol, Dr J. W. G. Lund, F.R.S., Freshwater
Biological Association, Ambleside, and Dr H. Belcher, Culture Collection of Algae and
Protozoa, Cambridge, provided valuable advice and discussion. Mr R. Ross, British Museum
(Nat. Hist.), kindly translated the Latin diagnoses.
 New algae from Antarctica 405

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BOURRELLY, P., 1968. Les algues d'eau douce, initiation ~ la systematique. Vol. 2. Les algues
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BROADY, P. A. The terrestrial algae of Signy Island, South Orkney Islands. British ,4ntarctic
Survey Scientific Reports (in press).
CHADEFAUD,M., 1943. Sur une sorte de "Fleur d'eau" form,~e par des Xanthophyceae du g~
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ForT, B., 1974. Taxonomie der Palmelloiden Chlorococcales (Familie Palmogloeaceae).
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PASCHER, A., 1932. l]ber einige neue oder kritische Heterokonten. Arch. Protistenk., 76: 1-82,
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zoospore-producing genera of the Chlorococcales. Indiana Univ. Pubis Sci. Ser. No. 20.
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