American Journal of Plant Sciences, 2011, 2, 381-395
doi:10.4236/ajps.2011.23044 Published Online September 2011 (http://www.SciRP.org/journal/ajps)
381
Vascular Plants of the Maritime Antarctic: Origin
and Adaptation
Ivan Parnikoza1, Iryna Kozeretska2, Viktor Kunakh1
1Institute of Molecular Biology and Genetics of National Academy of Sciences of Ukraine, Kyiv, Ukraine; 2Taras Shevchenko
National University of Kyiv, Ukraine.
Received April 7th, 2011; revised May 24th, 2011; accepted August 15th, 2011.
ABSTRACT
The question of why only two species of vascular plant have colonized Antarctica has not been fully answered. This
review is based on a series of parallel analyses of distribution, ecology, and adaptation on the morphological, cellular,
and molecular genetic levels, and addresses the causes of the exclusive adaptation of Deschampsia antarctica Desv.
(Poaceae) and Colobanthus quitensis (Kunth) Bartl. The authors conclude that the unique distribution of these species,
including the Antarctic Peninsula, is not related to the presence of any specific mechanisms of adaptation to the ex-
treme environment, but rather is a result of a gradual adaptation of these taxa to the extreme conditions during the de-
velopment of glacial events and wide distribution and a substantial seed bank which could ensure mosaic survival in
some ice-free areas, as well as survival through several years of snow and ice cover. Glaciological, molecular, popula-
tion and reproduction biology studies are still necessary to deepen our understanding of the timing of the colonization
of the region by vascular plants. However, keeping in mind that molecular methods alone are unlikely to give exhaus-
tive evidence, application of other adequate methods in the context of the history of Pleistocenic glaciation in the region
is also necessary to answer the question.
Keywords: Deschampsia antarctica, Colobanthus quitensis, Unique Adaptation, Time of Colonization, Gradual
Adaptation
1. Introduction
Antarctica, due to its geographical separation from other
continents and the presence of the oceanic polar frontal
zone and the Antarctic Circumpolar Current, is an iso-
lated continent [1-3]. The part of the continent where the
monthly mean temperature in the summer rises above
zero is separated into a distinct zone―the maritime Ant-
arctic [1,4]. Whilst the majority of the Antarctic conti-
nent is covered by permanent ice and snow, only 2% of
the landmass is available for colonization by plants and
animals [5]. The maritime Antarctic ecosystems host two
species of flowering plants: the Antarctic hairgrass (De-
schampsia antarctica Desv., Poaceae) and the Antarctic
pearlwort (Colobanthus quitensis Kunth. Bartl., Caryo-
phyllaceae) [6]. It is generally assumed that these species
have colonized this region by means of long distance
dispersal during the Holocene [1,7,8]. However, the fact
that these species are the only flowering plants present in
the native flora of this region raises the question of why
no other species characteristic of the southern sub-polar
regions have colonized the maritime Antarctic within the
period since the last glacial maximum [7,9], as a wide
range of other vascular plant species are known to be
present in the neighboring sub-Antarctic [7,70]. The same
latitudes in the Arctic, with its equally inclement habitats,
host a much larger list of vascular plant species [11].
A further complication is added by the recent rapid
warming periods in the region over the last 50 years [12].
This warming has equaled the most rapid rates globally,
demonstrating a rise of 3˚C annually along the western
coast of the Antarctic Peninsula [3,13-17]. Nonetheless, a
dispersal of other species adapted to polar climate has not
happened [7,9]. Only expansive weeds such as Poa an-
nua L. and P. pratensis L. that lack any signs of appro-
priate adaptation to polar environments have been able to
afford human-mediated colonization of this region [9].
However, no considerable expansion beyond the zone of
primary invasion of the only live population of P. annua
in the region has happened since more than 20 years ago
[1,18,19]. Reports about other species encounters in An-
tarctica appear from time to time, e. g. the finding of Na-
ssauvia magellanica J. F. Gmelin (Asteraceae) of Patago-

Vascular Plants of the Maritime Antarctic: Origin and Adaptation
382
nian origin on Deception Island (Downie, personal com-
munication, 2009). However, as a rule, such colonists are
not known to survive for longer than one season [1,7].
Interestingly, experiments with artificial transplantation
of the Patagonian and sub-Antarctic species to the region
have not been successful [7,9,20].
Profound adaptation to Antarctic conditions, depend-
ence on mycorhizas, as well as notable isolation, have
been put forth as the possible reasons for the lack of
vascular plants in this region [7,9]. However, the ques-
tion has not been answered yet and, in this review, we
analyse a body of evidence available across the bio-logi-
cal disciplines, in order to clarify the existence and the
importance of the specific traits of D. antarctica and C.
quitensis that ensure their survival and apparent success
in this region.
2. Systematics and Areal
The genus Deschampsia comprises 30 to 40 species dis-
tributed both in the Northern and the Southern Hemi-
spheres, the majority of which are perennials, though
there is a group of annual species as well [21-24]. Annu-
als from the Alps are supposed to have evolved from
perennial ancestors which had migrated from the plains.
The genus belongs to the Tribe Avenae, which is very
diverse in the Northern Hemisphere, although the genus
Deschampsia is more diverse in the Southern [22].
Taxonomic studies have shown that it is necessary to
separate the genus Avenella from Deschampsia [22,25].
Recent molecular phylogenetic studies have concluded
that D. antarctica from South America, sub-Antarctica
and the maritime Antarctic is closely related to the spe-
cies from southern Argentina and Chile: D. parvula
(Hook. f.) Desv. and D. venistula Parodi. Another closely
related branch comprises D. laxa Phil., D. kinga (Hook.
f.) Desv., D. berteroana F. Meigen from central Chile, D.
tenella Petrie from New Zealand, and the South Ameri-
can races of D. caespitosa (L.) P. Beauv. [21,22]. Our
study species—D. antarctica—is morphologically dif-
ferent from the other species of the genus from the sub-
Antarctic and the Andes, namely D. caespitosa, D. chap-
mani Petrie, and D. penicellata n. sp.; the differences lie
in the size of the stem, leaves, and the flower parts [26].
The distribution area of D. antarctica encompasses
Argentina, Chile and Peru. The species is also found on
Tierra del Fuego and the surrounding islands, the Falk-
land Islands, South Georgia, the South Orkney Islands,
the South Shetland Islands. It is also present on one of
the islands of the South Sandwich Islands archipelago, as
well as along the western coasts of the Antarctic Penin-
sula and the adjacent archipelagos of the maritime Ant-
arctic, reaching to the south Lazarev Bay on Alexander
Island [26; Convey, personal communication, 2008].
The genus Colobanthus Bartl. is distributed mainly in
the Southern Hemisphere (only C. quitensis is listed in
Mexico). Different authors give different numbers of
species in the genus Colobanthus [26]. Thus, for South
America the list varies up to 13 species, part of which are
now included in Colobanthus quitensis and the rest in C.
subulatus (D’Urv.) Hook. f. Of all the species of the ge-
nus, only C. quitensis is found in the maritime Antarctic,
again reaching Lazarev Bay on Alexander Island [Con-
vey, personal communication, 2008]. C. subulatus is
dispersed to the south up to South Georgia [26]. Overall,
the genus consists of up to 20 species [27].
Other species of the genus that are related to C. quit-
ensis are found on Kerguelen with the adjoining islands,
as well as on Heard Island―C. kerguelensis Hook. f.,
Tasmania, New Zealand, sub-Antarctic islands of the
Australian sector―C. muscoides Hook. f., Macquarie
Island―C apetalus (Labill.) Druce., as well as on the
Falkland Islands, Tierra del Fuego, and Patagonia up to
the latitude of 52˚25' S―C. subulatus [26,28]. C. quiten-
sis differs from these species in a number of morpho-
logical traits: the character of leaf tips, the leaf width, the
relative length of the sepal and the seed capsule, and the
sepal count [26].
C. quitensis and D. antarctica are characterized by
significant interpopulation variability of the traits men-
tioned above, which was the reason to separate the spe-
cies into a set of derivative species by early studies. The
distribution area of C. quitensis encompasses Mexico, the
highland regions of Ecuador, Bolivia, Chile, and Peru. It
is also present on Tierra del Fuego, the Falkland Islands,
South Georgia, the South Orkney Islands, the South She-
tland Islands, as well as along the west coast of the Ant-
arctic Peninsula with the adjacent archipelagos [26].
In the maritime Antarctic, along the west coast of the
Antarctic Peninsula, and on the adjacent islands the dis-
tribution density of the populations of both species of
vascular plants is heterogeneous. Their populations are
primarily located in the following three regions: 1) the
South Shetland Islands; 2) in the region between Cierva
Point and Cape Garcia; and 3) near Marguerite Bay. De-
tailed maps of the distribution of both species in the re-
gion have been provided by a number of authors [20,
29-33].
3. Ecology and reproduction
The habitat of the vascular plants and the plant commu-
nities they create is largely defined by the climate of the
region. As a result of the influence of the circumpolar
current a branch of which creates a buffering effect by
alleviating the temperature fluctuations in the region, the
climate of this part of Antarctica is oceanic [1]. Never-
theless, January is the only month when the mean air

Vascular Plants of the Maritime Antarctic: Origin and Adaptation383
temperature rises above zero. At the same time, the air
temperature rarely drops below –15˚C during the coldest
months (June through September). In this region, the
negative temperatures may occur at any time throughout
the year. However, even slight declensions in the land-
scape may play a crucial role in creating a microclimate
due to the low position of the sun in Polar zone. These and
other specific traits cause a variegated mosaic of mic-
roclimates, often differing even between the neighboring
islands [10].
Vascular plant vegetation areas are represented by rocky
slopes, moraines, simple soils (including peats) and peb-
ble beaches free of ice and the summer snow cover. At
smaller scales, plants can root on cliff ledges and in
crevices [20,29-31,34-37].
Both species appear not to be very demanding with
regards to the Antarctic soil characteristics. Nevertheless,
the slow organics decomposition processes driven by
invertebrates and fungi that take place in this substrate
still allow drawing an analogy line between this substrate
and soils [38]. Based on the main source of organics in-
flow, maritime Antarctic soils can be divided into two
basic groups: those formed under the Antarctic plant
communities [39] and ornithogenic. The latter are formed
in barren bird colony areas and are characterized by high
organics content [40].
Regarding the organics content in the soil, both vascu-
lar plant species of the region inhabit locations within
very wide ecological amplitude. This equally applies to
the content of microelements and trace metals [41-43].
Vascular plants develop a special Antarctic herb tun-
dra formation which comprises a single grass and tuft
chamaephyte sub-formation [6,10].
The plant starts tillering out early, producing shoots.
Young shoots are contained inside the leaf sheath. Leaves
are sessile, linear. The plant has bisexual flowers gathered
into tight acervuli. The species is considered self-pollina-
ting. Its flowers remain closed, so that self-pollination
makes for cleistogamy [44]. However, based on up-to-
date data, the possibility of cross-pollination can not be
completely excluded, as in South America both cleisto-
and chasmogamic flowers have been shown for this spe-
cies [45]. Chasmogamy may sometimes occur during the
mild seasons in Antarctica as well, as the ratio between
forming cleistogamic and chasmogamic (capable of cross-
pollination) flowers in other plant species has been
shown to depend on the environmental conditions [46,47].
In the Antarctic conditions, the development of D. ant-
arctica begins in November when seed germination and
recovery of last-year tufts starts [45].
The species is capable of vegetative propagation by
means of tuft outgrowth and split-off of the tuft parts. D.
antarctica plants often form a single dense and contigu-
ous tuft, which area varies from one to several hundred
square meters. A die-off of the plants in the central parts
of the tuft has also been documented. An uprooted plant
is capable of re-establishing after being transported to an-
other appropriate place [48]. As a result of this ability, the
possibility of plant dispersal by birds (as a nest building
material) has repeatedly been suggested [7,13,48].
C. quitensis is a perennial flowering plant which forms
dense low rounded hemispheric tufts and lives up to 35 -
40 years. This plant has a tap root, is almost incapable of
vegetative reproduction [49] and has an age structure
similar to that of D. antarctica [37,50]. The species, in
most if not all cases, is a self-pollinator. Its stamens ar-
ranged in front of the pistil make for a high probability of
cleistogamy, which leads to inbreeding. The seed yield
resulting from one self-pollination event is rather high:
near 43 seeds per capsule in plants from Tierra del Fuego
and the Ands [26].
There is almost no evidence of vegetative reproduction
in C. quitensis, and all studied populations seem to have
originated from seeds. Under favorable conditions, seeds
of this species can survive for a long time. The germi-
nation temperature may be high. Seed germination disrup-
tion in C. quitensis has been observed when the tempera-
ture dropped from 9˚C to 2˚C [49].
In C. quitensis, just like in other perennials with slow
growth in extreme environments, the appearance of new
plants from seeds occurs only in some years [49]. None-
theless, this is the groundwork of its population renewal
[11]. Studies of the population structure of C. quitensis
have revealed its extreme irregularity. Low reproduction
rates are common for Antarctic and highland plants, as
the success of seed reproduction and seedling survival
are severely restricted by the unfavorable climatic factors.
It has been shown that on some islands the reproductive
success of C. quitensis depends on the conditions during
a particular year. This species blossoms irregularly on the
Argentine Islands and in other places of the maritime An-
tarctic. In some years, high seedling mortality has been
registered. A successful seed reproduction is rare in such
species, being almost exceptional [51]. Thus, it has been
demonstrated for nine Arctic herb species that the pro-
portion of flowering plants varies from year to year in the
range between 0 and 40%, and the percent of surviving
seedlings—between 0 and 96% [49].
In the case of the species discussed, dispersal of tufts
by birds may also be possible. The modelling of this kind
of dispersal of C. quitensis tufts that the authors carried
out in different zones of Point-Thomas Oasis demon-
strated that their successful rooting is possible on rela-
tively wet areas of the ice-neighboring zone [48]. An
increased fitness of both species under the dynamic con-
ditions of the Antarctic environment may also be pro-

Vascular Plants of the Maritime Antarctic: Origin and Adaptation
384
moted by their ability to form seed banks yet described
within the region [52]. Meanwhile, newcomer species
can not generate a sufficiently large seed bank, and con-
sequently a sufficient number of populations, in time,
which doesn’t favor their establishment and may cause
early elimination, just like it happened, for instance, to a
population of Poa annua that had survived through 13
years and was obliterated in 1967 by a volcano eruption
on Deception Island [9].
Development of mycorrhiza cannot be considered as
unique adaptation factor for either of the species as is not
a specific process, and the fungi revealed are not unique
to these vascular plants due to their vast distribution in
the nearby sub-Antarctic [53].
Overall, the life forms, ontogeny, and reproduction of
the Antarctic vascular plants do not show any radical
distinction from other closely related or having the same
life-form species of the Polar Regions or the highlands
[11,54], whose reproductive biology is well adapted to
the inclement environment of their habitat.
4. Anatomical and Biochemical Adaptation
The investigation of both Antarctic flowering plant spe-
cies has not revealed any traits that could allow for a
qualitative distinction from the other polar species and
would explain the better survival of these species in the
most inclement regions on Earth [55-57]. In general,
their anatomy is characteristic of the plants inhabiting
arid places. Leaves have stomata and their upper sides
are coated with a thick layer of wax, which is one of the
traits representative of the drought-resistant plants. The
hairgrass is known to have a wide range of individual
forms which differ by the chloroplast count in the cells of
the inner leaf tissue―mesophyll. The number of chloro-
plasts negatively correlates with such parameters of the
environment as short daylight hours and low average
annual temperatures [57].
Peculiar traits of the photosynthetic system of both spe-
cies have been a separate study topic. In general, the
photosynthetic system of vascular plants is well adapted
to work under low temperatures. However in these Ant-
arctic species it becomes photosynthetically inactive
when the temperature drops below –2˚C, just like in all
other vascular plants [9,58].
The search for the reasons of this uniqueness was di-
rected towards the biochemical adaptations to living un-
der low temperatures and severe UV-exposure conditions
[59]. In doing this, the majority of researchers addressed
D. antarctica, and to a much lesser degree C. quitensis.
On the biochemical level, D. antarctica has a system of
adaptations typical, to a varying extent, of all cold habitat
plants. There is a group of stabilizer hydrophylic pro-
teins―dehydrines―the intensity of synthesis of which
alters under a low temperature stress [60] and for which
there are several genes identified in the hairgrass. One
part of the products of these genes accumulates under an
external influence by abscisic acid (ABA), and another
part―under osmotic and salt stress, which is demon-
strated by the presence of an ABA-dependent and ABA-
independent pathways of dehydrine synthesis regulation.
Analysis of the pool of these proteins has identified
seven stress proteins accumulating under the low tem-
perature stress in vascular and tectorial tissues where the
zones of initial ice formation are found [61].
It has been shown that heat shock proteins (70 kDa)
also accumulate in D. antarctica under the temperature
stress and may well explain its low photosynthetic opti-
mum temperature (+13˚C) [5,62].
Tests for the presence of anti-freeze proteins typical
for plants (irrespective of the area of their distribution) in
D. antarctica have revealed their proportion in the gen-
eral protein pool to be rather high [63]. Additionally, an
IRIPs (recrystallization inhibition proteins) gene has also
been identified in D. antarctica which codes for a protein
that inhibits water recrystallization in the extracellular
space. However, the protein is not species-specific [64].
Sequences have been identified in the D. antarctica
genome which are homologous to the sequence of the
genes coding for ubiquitin-like proteins. In plants, these
proteins participate in the ubiquitin-ATP-dependent pro-
tein degradation and, in particular, prevent self-fertiliza-
tion and are involved in reactions to stress [65]. Under
low temperatures, activation of antioxidant enzymes [66]
and soluble carbohydrate accumulation in tissues [59]
have been revealed as well. A gene has been studied in D.
antarctica which codes for the saccharophosphatesyn-
thetase enzyme. The activity of the enzyme is known to
increase in response to low temperature, although its qua-
ntity and the expression of the gene remain stable [67].
Investigation of the lipid composition of the D. ant-
arctica membranes has not revealed any special lipids.
However the phosphatidyl glycerol content is lower, which
is commonly linked to higher sensitivity to stress [68]. A
comparison between the pigment-protein complexes of
the thylacoid membranes of D. antarctica and Pisum
sativum L. [69] has not revealed any differences either.
At the same time, quantitative differences have been found
in the general content of different pigment-protein com-
plexes [70]. D. antarctica’s defence against UV-expo-
sure is activated by means of an increase in β-carotene
content and a reduction of violaxanthin, as shown for the
UV-treated leaves. The role of carotenoids may be linked
to the defensive increase in thylacoid membrane fluidity
as a reaction to damage produced by oxygen upon expo-
sure [71]. An increase in flavonoid content has also been
shown as a mechanism of defence against UV-exposure,

Vascular Plants of the Maritime Antarctic: Origin and Adaptation385
however this type of reaction is not specific to Antarctic
plants [72].
Gene database search (GenBank, TrEMBL, and Swiss-
Prot (http://www.expasy.org; Supplement 2) revealed 37
amino acid sequences that can be determined which are
part of the D. antarctica proteome. Of these, 20 sequen-
ces are potential proteins or protein fragments with a
predicted function, and 17 sequences are potential pro-
teins or protein fragments whose function has been de-
termined. It has been shown for D. antarctica that the
transcripts of 25 genes previously described for other
species appear as a reaction to cold [73]. The analysis
performed shows that the annotation of the genome and
the proteome of D. antarctica is in its initial stage, and
the functions of most known potential proteins have yet
to be determined [70,73].
5. Cytogenetic Traits
The total chromosome number of the species of the ge-
nus Deschampsia is usually 2n = 26, with the basic count
X = 13. There are only some species with the basic
chromosome count 7, such as D. artropurpurea (2n = 14)
and D. flexuosa (commonly 2n = 28), which, according
to molecular taxonomy data, are isolated as separate
genera [22,24,74] (Table 1).
Differences between the Deschampsia core and D. at-
ropurpurea and D. flexuosa have been demonstrated by
studying isozyme spectra, C-banding, and using methods
of plastid and nuclear DNA restriction patterns [22,24].
Analysis of the data in Table 1 reveals that irrespec-
tive of the areal (and in the majority of cases the species
of the genus are adapted to living in cold wet meadow
habitats) speciations were not accompanied by changes
in chromosome numbers. However, polyploidization and
aneuploidization of the genome have been found. Karyo-
logical variations in the species of the genus Deschamp-
sia, namely D. caespitosa, are caused by the ability of
smaller chromosomes to merge with subsequent poly-
ploidization [22,77]. Based on this, Poaceae have been
attributed traits of ecological differentiation linked to
ploidy, with diploid plants having lower degrees of po-
tential realization of their niche than polyploids. The de-
gree of this realization increased with an increase in
ploidy [78]. It seems like it is this trend that lies at the
mainstay of the appearance of new species forms that are
known to have only tetraploid (with n = 13) karyotype,
such as for instance D. obensis, D. mackenziana, and D.
mildbraedii (Table 1).
The high proportion of aneuploids and the variation of
the diploid chromosome count from 18 to 26 have also
been shown by cytological analysis for D. caespitosa
from northern Lake Ontario populations (Canada). Be-
sides, individuals with 2n = 26 are known to contain ad-
ditional so-called B chromosomes. The role of the latter,
also identified in the genome of D. wibeliana, remains
unclear [75,79]. In some specific habitats polyploidiza-
tion has been found to lead to isolation of endemic forms
[76].
Regarding D. antarctica, its karyotype, according to
the only data available up to date [24], is 2n = 2x = 26,
with the karyotype formula 10 m + 6 cm + 2 t. The nu-
cleolar organizer region is located on the short arm of
one of the submetacentric chromosome pairs which form
a terminal satellite. The authors have also detected ane-
uploidy (aneusomy). The investigation of the secondary
roots in D. antarctica from Galindez, Petermann, and
Berthelot islands (the location of Ukrainian Vernadsky
station, the maritime Antarctica) elucidated the whole
picture. Additionally, polysomy has also been revealed.
The chromosome count variation in root radix meristem
cells was rather high―from 10 to 68 chromosomes.
Therefore, D. antarctica from this region not only dem-
onstrates polysomy with a range of variability of the
chromosome count comparable to that of the genus, but
is also characterized by frequent aneuploidy [80].
It is generally believed that species with low DNA
content are better adapted to lower temperatures, i.e. DNA
content may be considered as one of the factors that in-
fluence geographical distribution [7]. D. antarctica’s low
DNA content (10 pg) does indeed characterize the spe-
cies as one preadapted to develop in cold habitats [81].
A reaction of the plant’s leaf tissue interphase cells to
alterations in environmental conditions has been dem-
onstrated. Analysis of DNA content in the nuclei of leaf
parenchymal and epidermal cells in plants from differ-
ent populations from the Argentine Islands and the
Point Thomas oasis has revealed statistically significant
differences of this parameter in plants from different
places. Therefore, with regards to nuclear DNA content
(the degree of ploidy) the genus Deschamp sia, includ-
ing D. antarctica, is a complex heterogenous entity.
Ploidy and DNA content vary significantly, which, in our
opinion, is of adaptive importance [42]. This is congruent
with the ideas about the mechanisms and causes of these
phenomena proposed by other authors who link the ob-
served aneusomy to the differential influence of envi-
ronmental factors, temperature in particular (see discus-
sion in [24]).
Regarding C. quitensis, its 2n chromosome set in-
cludes 80 chromosomes. Due to the small size of the
chromosomes, karyotype investigation has not been com-
pleted so far. Near 80 chromosomes have been found in
other close species, such as C. apetalus and C. affinus
[26].

Vascular Plants of the Maritime Antarctic: Origin and Adaptation
386
Table 1. Chromosome numbers of Deschampsia genus specie s, and its gener al distribution and ecological grow conditions [24,
74-76].
Specie Chromosome
number Distribution Ecological grow conditions
D. caespitosa = D. refracta = D.
media 26(0-2В), 26, 52 Cosmopolite Stream valleys
Species, according to other data subspecies D. caespitosa
D. alpina
26-52, (38-39),
39, 41, 49, 50,
52, 56
Europe, inter alia circumpolar regionsHumid grasslands and rocky substrates
D. bottnica 26 Bothnia, Bothnia Bay Humid grasslands and rocky substrates
Deschampsia obensis 52 Eastern Siberia, Russia Humid grasslands and rocky substrates
D. mackenziana 52 Canada, especially Athabasca Sandy biotopes
D. glauca 26, 52
Eurasian Subarctic, circumpolar Ural,
Jamal, North (from Alaska and the
Yukon south to northern Mexico and
east to Montana, Wyoming, and
Arizona) and South America
(disjunction in Chile).
In tundra, on rocky and gravel-stone flood
plains near bogs, on cryophilic grasslands
on gravel soils, in moist to wet habitats,
from near sea level to alpine elevations
D. wibeliana 26(0-5В) Europe, Wadden Sea coast Wet meadows
D. rhenana 49-52 Europe Wet meadows
D. holciformis 26 North America: USA, Canada In coastal marshes and sandy soils
D. orientalis = D. sukatscewii = D.
borealis = D. brevifolia = D.
pumila = D. borealis = D.
paramushirensis
26, 28, 36, 52 Europe, and North America Arctic,
China, Japan
Wet meadows and other humid tundra
biotopes
D. festicaefolia 27, 28 Alps, Japan Alpine biotopes
D. beringensis 26, 42 Alaska, Canadian and Russian ArcticCoastal wet biotopes
D. paviflora 26, 28 Europe Wet meadows
D. macrothyrsa 26 Far East, Sakhalin Wet meadows
Other species
D. atropurpurea 14 Cosmopolite Large spectrum of grassland ecosystems
D. setaceae 14 Cosmopolite Large spectrum of grassland ecosystems
D. flexuosa 14, 26, 28, 32, 42Cosmopolite On dry, often rocky slopes, and in woods
and thickets, often in disturbed sites
D. liеbmanniana 26,52 Mexico Rocky alpine biotopes
D. tenella 26 New Zealand endemic Mountain biotopes
D. chapmanii = D. novae-zelandii 26, 28 New Zealand endemic Mountain biotopes
D. gracillima 26 New Zealand endemic Subalpine and alpine zones, wet rocky
slopes
D. danthonioides 26
Western part of North America
(California), South America (Andes,
Central Chile)
On dry slopes, perturbed places, gravels,
road sides, grows in temperate and
cool-temperate regions, usually in open, wet
to dry habitats and often in disturbed
ground.
D. elongata 26 Cosmopolite River banks, wetlands, mountain meadows
D. nubugenia = D. australis =
D. hawaiiensis 26 Hawaii Islands: endemic to Kauai, Molokai, Maui, Hawaii Islans Wet meadows
D. koelerioides 26
Asia, Siberia, Altai region, Central
and Western Asia, Mongolia,
described from Tien Shan
On alpine zone, grassland patches near
streams
D. komarovi 26, 52 Eurasia Meadow biotopes
D. mildbraedii 52 Africa West-central tropical part
D. antarctica 26 South America, sub-Antarctica,
maritime Antarctica
Antarctic tundra, grasslands of
sub-Antarctica, Patagonia and mountain
grasslands of Andes
D. pamirica 26 Pamir and other Asian highland
ridges Highland biotopes
D. argentea-maderensis 26 Madeira Meadow biotopes

Vascular Plants of the Maritime Antarctic: Origin and Adaptation387
6. Evolutionary and Phylogenetic Aspects
While recognizing D. antarctica and C. quitensis as Ant-
arctic indigenous species, the majority of researchers ad-
here to the hypothesis of post-Pleistocenic colonization
of the region by these plants [5,7,8,30]. At the same time,
the exclusive existence of only these two species and the
traits of their high gradual adaptation allow one to sug-
gest that their earlier colonization of the region hap-
pened during the Tertiary period [82]. Since D. antarc-
tica and C. quitensis are by no means invasive and have
yet not colonized a number of suitable habitats in the re-
gion (for instance C. quitensis are absent from all of
South Sandwich Islands, D. antarctica on majority of it
[16]), it seems reasonable to theorize that the expansive
species from the sub-Antarctic, rather than these two,
should have colonized the emerging suitable habitats
after the ice retreat during the post-Pleistocene transmis-
sion. However, provided these species have been early
colonists, not only their gradually adaptation but a simple
saturation of the Antarctic environment with their dias-
pores would have given them a selective advantage in
surviving the hardship of the Antarctic environment.
Concerning the age of these species on Earth in general,
and in Antarctica in particular, there are only scanty data.
The results of paleobotanical studies indicate that
Poaceae, just like Caryophyllaceae, appeared during the
late Cretaceous and had spread over the supercontinent
Gondwana before it completely separated in the Tertiary.
The first findings of grasses and their communities relate
to the South American Eocene (45 mya). Although pol-
len grains have been found in the African Eocene, grass
communities appear there only in the late Myocene (14
mya) [83]. The first finding of a pollen grain of Caryo-
phyllaceae—Caryophylloflora paleogenica G. J. Jord.
and Macphail—occurred in Australia and New Zealand
and relates to the late Cretaceous [84]. This species may
well have pertained to any other caryophyllid group sensu
lata [8], however true Caryophyllaceae themselves ap-
peared in South America not later than in Myocene [84].
In the case of Poaceae, in Myocene there was an expan-
sion of these C4 plants over the adjacent territories with a
formation of open grass communities [85].
Attempts to describe the directions and timing of the
general Antarctic vascular plant species dispersal have
been made only for Deschampsia. There are two hypo-
theses of the initial dispersal of the species of this genus:
the northern one, which assumes that the genus devel-
oped at high and middle latitudes of Eurasia and only
later spread to the south where a secondary speciation
center formed, and the southern one with the origin point
in South America, which assumes the genus to be very
old in the Southern hemisphere [21]. The common clade
for South American species together with D. antarctica
(batched based on molecular genetics data) corroborates
the localization of the species creation center of at least
this part of the genus near the points of contact between
South America and Australia and New Zealand via the
Antarctic, which may have been the case all the way up
to Pleiocene [21,83]. Based on the general idea by
Hooker who admitted in 1851 that modern southern flora
might represent remnants of the flora of Gondwana [1], a
concept has been coined that D. antarctica and C. quit-
ensis appeared in the region before Pleistocene. In view
of the contact between South America and Antarctica via
the chain of the Scotia Islands, dispersal all the way up to
the late Tertiary when the Antarctic ice sheet still didn’t
cover the whole continent can not be excluded [82]. At
the same time, the vicissitude of glacial maxima and
minima that took place during that period in the maritime
Antarctic [86] didn’t lead to vegetation extinction, and so
might concurrently make for its gradually adaptation
(mainly thereby wide distribution and dence seedbank) at
the onset of the Pleistocenic maximum. In a reply to this
suggestion, it was noted that in case of such an ancient
age of these species they should have gone through a
significant divergence leading to new younger species
and subspecies [8]. Perhaps, one could look for the traces
of such a divergence in the previously described close
relatedness of the sub-Antarctic species of the genus
Colobanthus, as well as in the attempts to isolate 13 spe-
cies within the genus Colobanthus instead of the two—C.
quitensis and C. subulatus—from South America [26].
Contrary to what many researchers might expect, such a
variability of closely related forms interpreted as separate
species might not gain speed to develop further due to
the narrow range of ecological constraints a form was ob-
liged to fit in to have a chance to survive in the inclement
environment of the Andean highlands or Antarctica grip-
ped by ice.
Still, there remains an open question about the possi-
bility of vascular plants’ survival in the region through
the glaciation events during Pleiocene—Pleistocene (20 -
1 mya). Information on the scale of glaciation in the
Southern hemisphere would be of much help in answer-
ing this question. However, determination of ice bounda-
ries is often complicated due to their erasure or masking
by later events [3]. Based on data by Sugden and Clap-
perton [87] and Law and Burstall [28], Smith notes that
Antarctic vegetation could hardly remain to the south of
South Georgia and Heard Islands which probably were
fully covered with ice [1]. At the same time, a reference
to a map of glaciations on the Antarctic Peninsula during
the Pleistocenic maximum compiled based on a whole
series of publications suggests that the South Shetland,
South Orkney, and South Sandwich Islands were outside

Vascular Plants of the Maritime Antarctic: Origin and Adaptation
388
the zone of contiguous ice sheet [88]. Also, although
Tatur [89] with a reference to Clapperton [90] presents a
map illustrating the total glaciation of King George Is-
land, it still contains areas marked as free of ice. Marsz
[91] points out a series of regions on this island which,
due to a number of reasons, could not freeze over.
In such conditions, the possibility of the ice-free zones—
Antarctic refugia—remaining in the region should be
considered [82]. For instance, two territories that have
not been covered with ice and a number of invertebrate
species that have survived have been demonstrated [3,92,
93]. However, in the case of vascular plants of the mari-
time Antarctic, only the areas close to the ocean which
are exposed to the buffering effect of the ocean could
probably serve as refugia, as it is this effect that allevi-
ates the climate oscillation [1]. Due to the diverse relief,
exposition, and the underlying rock composition, a vari-
ety of microclimates form, some of which may be suit-
able for D. antarctica and C. quitensis. This seems to be
the only plausible reason why the species are absent from
many maritime Antarctic shore areas under the condi-
tions of the progressing warming in the region. There is
also evidence that this, together with the geomorpho-
logical and other characteristics, hinders ice sheet forma-
tion on many maritime Antarctic territories [91]. Ac-
cording to Marsz, this kind of oases can be observed on
King George Island. One of them, the Point Thomas oa-
sis, is characterized by some of the largest tufts of Ant-
arctic herb tundra formations in the maritime Antarctic
[32,94]. The present day area of such oases compared to
that of bare vegetation-free morains is relatively small.
The oldest peat deposits on sub-Antarctic and Antarctic
Islands are known to be of Holocene age (5 - 6 thousand
years; [1,3]. However older deposits may have not re-
mained as a result of regular washouts or sliding of large
masses of peat into the ocean driven by ice streams and
the ice tongues of the many ice encroachments rather than
glacial rock overlay.
7. Molecular Genetics Data
As part of the discussion on the directions and timing of
the Antarctic vascular plants’ colonization, a reference to
the molecular genetics data should be included which,
according to some authors, suggest a relatively recent
colonization of the region [8]. Indeed, it is a generally
accepted idea that biota of the Antarctic refugia must
demonstrate a notable genetic diversity as a result of the
accumulation of mutations with their fixation by in-
breeding and the absence of gene drift [95]. On the con-
trary, recent colonists are believed not to have enough
time to generate interpopulation diversity and demon-
strate the founder effect.
A large amount of data on D. antarctica heterogeneity
has accumulated from research based on the AFLP
method. However, an important limitation of the method
is its inability to identify the genomic sequences respon-
sible for the detected heterogeneity. For this reason, in-
terpretation of the results obtained by AFLP may be
complicated [96]. The interpretation of the results of ear-
lier D. antarctica genetic polymorphism studies have
been ambiguous. A study using the AFLP method has
demonstrated, according to the authors’ interpretation, a
low variability—13% between populations from Signy
Island (the South Orkney Islands) and Anchorage, La-
goon, and Leon Islands that are 1350 km away from the
first one. Nevertheless, AFLP method results inevitably
bring about the question of what is the actual age of the
divergence a given heterogeneity accounts for. At the
same time, a high polymorphism has been revealed be-
tween the populations from different parts of Signy Is-
land, and a low one between those from the southern
region. Additionally, the absence of identical genotypes
was registered in both regions [97].
Chwedorzewska, also based on AFLP, has found a
higher interpopulation heterogeneity in D. antarctica
compared to that of the Arctic species D. brevifolia R. Br.
and D. alpina (L.) Roem and Schult. from the Svalbard
archipelago, with the latter two species demonstrating
clear evidence of being post-Pleistocenic colonists. At the
same time, D. antarctica populations of maritime Ant-
arctic farther south demonstrated lower heterogeneity
than northern populations living in a less hostile environ-
ment [98]. Such a pattern, apparently, can be explained
by a stepwise dispersal of the species in the region [8].
Concerning the possibility of multiple colonization events,
provided several genotypes have remained in different
Antarctic oases, their counter-dispersal during the period
of milder climate conditions on from the beginning of
Holocene may well have made for the pattern obtained
by Chwedorzewska. This explanation is corroborated by
another two studies on interpopulation polymorphism in
D. antarctica, also using AFLP. The co-existance of the
two groups of genotypes and intermixed populations has
been revealed in a study of the molecular heterogeneity
in populations from the Point Thomas oasis (King
George Island) [99].
Additionally it has been shown that interpopulation
heterogeneity in plants from the Falkland Islands is
higher than that between other maritime Antarctic popu-
lations that live farther apart from each other. On the
other hand, an analysis of heterogeneity in D. antarctica
from the far more distant South Shetland Islands and
Argentine Islands has revealed that the plants from both
regions are equally heterogeneous, which didn’t allow
their clear-cut clustering with further batching them in
agreement with their geographic location [100]. The

Vascular Plants of the Maritime Antarctic: Origin and Adaptation389
higher heterogeneity in northern regions, again, provides
evidence of more genetic variants that have formed here,
while the heterogeneity level-off in populations from
regions farther south may be explained by a northern
origin of these populations, which is in accordance with
the previous study.
Therefore, AFLP data can suggest the localization of
the center of genetic diversity of D. antarctica on the
north of the maritime Antarctic [97] and the adjacent
archipelagos of sub-Antarctic, which, in our opinion, is
in good agreement with the possibility that refugia have
existed here in which the species might survive glaci-
ations. Such a picture, in the case of Antarctic vascular
plants, could well make use of the Wladislaw Szafer’s
idea of migration relicts as species relict only in some
parts of the areal, i.e. those that dispersed from refugia
into adjoining regions at a later time [82,101]. In the case
of Antarctic plants, survival may have been successful in
a number of refugia close to the sea on the South Shet-
land Islands and the South Orkney Islands, as well as,
perhaps, the South Sandwich Islands from which indi-
vidual plants with relatively heterogeneous (after isola-
tion) genotypes might counter-disperse into both the
more southern territories of the maritime Antarctic and
the nearest ice-free regions. Preservation of glacial refu-
gia on South Georgia Island has also been suggested
[102]. The presence of flowers capable of cross-pollina-
tion in D. antarctica indicates that gene transfer between
the usually cleistogamic populations may also be possi-
ble during particularly favorable seasons.
Such data are absent for the second Antarctic vascular
plant species, however the notable morphological vari-
ability of C. quitensis, as well as the concentration of
species of this genus in the northern parts of the maritime
Antarctic, Scotia and the southern end of South America
[26] is in good agreement with the idea of the existence
of refugia specifically in this region.
More specific conclusions based on the available body
of AFLP data might be expected if it included popula-
tions from South America. Investigation of these mater-
nal (apparently in both cases—pre-Pleistocenic and post-
Pleistocenic colonization) locations would probably help
to clarify the time of the divergence. However in this
case, one should keep in mind that those Andean glaci-
ations, similar to those in Antarctica, significantly re-
stricted the territories suitable for vegetation develop-
ment. As a result, the territories that became free of ice
after the glaciation periods could have been colonized by
plants from both the neighboring populations and those
transmitted from the regions farther south.
One of the reasons why AFLP data should be ap-
proached with a great deal of caution becomes apparent
from the studies on genetic heterogeneity in a population
of the annual grass Poa annua L., a species which is
known to be a transferred weed that appeared near the
Polish station Arctowski in the 1980s. Admitting that the
station personnel is accountable for transferring the plant
from Europe, in contrast to the expected low heterogene-
ity in this species as a result of the founder effect,
Chwedorzewska points out that the actual heterogeneity
is surprisingly high—60% [103]
A concurrent study of D. antarctica from South Ame-
rica, sub-Antarctic, and the maritime Antarctic has been
done only with respect to non-coding chloroplast DNA
[100]. Only three haplotypes of the chloroplast DNA
have been found, with heterogeneity obtained with only a
small fraction of primers. Additionally, very large terri-
tories were represented with just a few samples in this
study. The revealed unique for the South Orkney Islands
haplotype C within the zone of contiguous distribution of
haplotype A brings about the idea that plants with dif-
ferent genotypes can neighbor on the same island or a
close group of islands. Therefore, the existence of plants
bearing these and some as yet unknown haplotypes in
other regions can not be ruled out. Based on the revealed
haplotype C unique for the Orkney Islands, one could
suggest a possible refugium located here. However, there
is evidence of the putative glaciation of the whole area
south of 60˚ [87]. At the same time, the absence of such
evidence from the Indian Ocean side (the Kerguellen and
Crozet archipelagos) allows the authors to describe the
haplotype formed here as a result of the isolation in refu-
gia near the ice sheet edge [100].
In order to be able to come to safer conclusions about
the age of the divergence between species living in South
America, sub-Antarctic, and maritime Antarctic, some
researchers turned to employing molecular markers that
are more use-proven in phylogenetic plots. The authors
are aware of only one study in which several specimens
of С. quitensis from the Chilean Ands and the maritime
Antarctic were compared with regards to the internal
transcribed spacer (ITC) of ribosomal DNA. The vari-
ability of the nucleotide sequence of this, common in
taxonomy, region of DNA was only 1.17% [104].
However 35S rDNA (the nuclear locus coding for 5.8S,
18S and 25S rRNAs; for review see [105] represents a
class of repeated sequences under control of concerted
evolution which is responsible for the high degree of
homogenization between repeats [106]. The high degree
of homogenization and the existence of regions evolving
with different rates make 35S rDNA a very attractive tool
for molecular taxonomy, phylogeography, and popula-
tion genetics. In particular, comparison of rapidly evolv-
ing ITS1 and ITS2 has widely been used for taxonomic
reconstructions among members of the same or closely
related genera [107,108].

Vascular Plants of the Maritime Antarctic: Origin and Adaptation
390
For instance, Saxifraga paniculata Mitt., which has
undoubtedly survived highland glaciations in separate
refugia, demonstrates only 14 variable nucleotides (2.4%)
out of 583 nucleotides of ITC on the interpopulation
level [109]. In the Mediterranean plant genus Anthyllis,
the species Anthyllis montana L. is known to have evol-
ved in late Pleiocene through early Pleistocene. It differs
from its closest species by 5, 2, and 17 nucleotide substi-
tutions in ITC. Its intraspecific divergence started only in
late Quarternary (0.7 mya) [110]. A similar history, in
the case of D. antarctica and C. quitensis, could poten-
tially explain the absence of clear-cut species separation
for the significant geological time some authors propose
[8], as well as the onset of species split in the post-glaci-
ation period, which seems to have taken place in the re-
gion with C. quitensis. Interestingly, based on the data on
interpopulation variability of ITC in 12 plants of D. ant-
arctica from 6 sites in the region of the Argentine Islands
and 6 sites on King George Island (the South Shetland
Islands separated from the former group by 500 km), as
well as data available from GenBank (http://www.ex-
pasy.org), a high degree of identity (96.3%) between the
samples and GenBank data was observed with the total
differences between plants from different populations
being within several nucleotides [96]. Nonetheless, ana-
logy with the abovementioned Anthyllis montana is hard-
ly possible.
The 12 analyzed samples of D. antarctica from both
regions seem to have had different genetic origins. The
evolutionary ancestral ITS variant and a derived variant
have been found in both locations, whereas the most di-
vergent variant has only been detected on King George
Island. Therefore, these results demonstrate that geneti-
cally distinct plants may co-exist within the same or ad-
jacent populations on Antarctic islands [96]. Similar ob-
servations have recently been made for chloroplastic
DNA by [100]. Based on these data, a spread of plants
with different genotypes closing in on their way during
recolonization of post-glacial territories can be assumed.
Consequently, the molecular genetics studies carried out
on the population level for Antarctic vascular plants so
far do not allow us to unambiguously determine the time
of D. antarctica and C. quitensis colonization. In view of
this, a determination of the precise timing of such a
spread, as well as the age of the above-mentioned geno-
types, as yet seems problematic. The problems stem from
the uncertainties in the applied methods, as well as the
insufficiency of the available material, both the living
plant samples from the maritime Antarctic and the fossil
records. Additionally, it should be admitted that molecu-
lar biology studies of these species are so far at the be-
ginning, and this promises many unexpected data to ap-
pear in future.
8. Conclusions
The unique dispersal and appropriate adaptation of only
two species of vascular plants in the natural flora of the
Antarctic remains enigmatic. Possible approaches to
solve this problem were considered to lie in the domains
of the subcellular and molecular levels of organization of
plants from different populations of these species. Indeed,
complex studies of these aspects in Antarctic vascular
plants are only beginning. At the same time, the multi-
farious data available so far on these species provide
evidence that the causes of the success of Deschampsia
antarctica and Colobanthus quitensis are not related to
any unique adaptations but to their history of migration
and adaptation to living in the region. In this paper, it is
proposed that the explanation of this phenomenon may
lie in of a prolonged spreading of these plants in condi-
tions of gradual worsening in the maritime Antarctic with
the alternating development of newly formed ice-sheets
thereby a wide distribution and formation of a spatially
dispersed seed bank. Both factors could potentially have
allowed the two species to survive in sporadic areas that
were free of ice sequentially during but a few years. In
regard to this, further complex studies of glaciations and
microclimates that have been the case in the region dur-
ing a sequence of glacial events are expected to be the
most tempting perspective in the context of the history of
the regional flora. And it is only approach employing
molecular genetics, population and reproductive biology
studies that will be able to explain the high gradually
adaptation of these species. A study of the genome evo-
lution of both of the Antarctic genera from different re-
gions of their areal using adequate molecular genetics
and other methods is one of the main steps in the future
research. At the same time, it would be informative to
compare the obtained data with that for the Arctic species
of the genus Deschampsia and the genera close to the
genus Colobanthus.
9. Acknowledgements
We thank anonymous reviewer, A. Rozhok & M. Rozhok
and P. Convey for their friendly help with manuscript
preparation and English correction.
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