Pugnaire F.I. Valladares F. Functional Plant Ecology
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Some estimates of annual production are given in Table 13.5. There is also little difference
between the higher rates for both lichens and mosses and both show a strong decline from the
maritime to the higher latitudes. The extreme climate conditions of the continent depress
production, which suggests that there is little adaptation by the plants. It is unfortunate that
so many estimates come from the maritime although, in defense, it is one of the major
cryptogam-dominated areas of the world. Overall, however, the data are very thin and
more and better measurements are needed, especially in the continental Antarctic.
GROWTH RATES
It is difficult to be sure just how reliable the productivity data are because of the assumptions
involved. One check is to compare what is known about growth rates in the same areas.
TABLE 13.5
Estimates of Annual Production for Higher Plants, Lichens and Mosses at Several
Locations in Antarctica
Species Location
Annual Production
mg gd:wt:
1
(1)
gm
2
y
1
(2)
mg CO
2
gd:wt:
1
(3) Comments
Higher plants
Deschampsia
antarctica
Signy Island
(608 43’’ S)
1700 (2) 390 (2) Estimated from model based on
microclimate and NP data. Gravimetric
(Edwards and Smith 1988; Edwards 1972)
Lichens
Cladonia rangifera
Usnea antarctica,
Usnea aurantiaco-
atra
Signy Island 80–200 (1) Gravimetric (Hooker 1980)
U. antarctica King George
Island
(628 09
0
S)
<300 (3) Estimated from model and year-round
microclimate studies (Schroeter et al.
1995)
U. aurantiaco-atra Signy Island
Livingston
Island
(628 40
0
S)
250 (2) 85 (3) Smith (1984), Schroeter (1997)
Usnea sphacelata Casey
(668 17
0
S)
14 (1) Estimated from model and assumed season
length (Kappen et al. 1991)
Cryptoendolithic
lichens
Dry Valleys
(778 35
0
S)
3 (2) Estimated from NP data and microclimate
(Kappen 1993a)
Mosses
Chorisodontium
Polytrichum spp.
Signy Island 315–660 (2) Harvest plus prediction (Collins 1973, 1977,
Longton 1970)
Calliergidium þ
Drepanocladus
spp.
Signy Island 223–893 (2) Davis (1983)
Bryum
pseudotriquetrum
Syowa
(698 00
0
S)
16 to þ13
(mean þ3.7)
Modelling from microclimate (Ino 1983)
Bryum argenteum Ross Island
(778 50
0
S)
100 (2) Modelling from microclimate þ NP
reponses (Longton 1974)
Note: The sources of the data are given in the comments column. The numbers in brackets in the annual production
column refer to the units for the production.
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420 Functional Plant Ecology
Examples are given in Table 13.6 from a variety of sources. Once again the vast majority of
estimates are from the northern maritime zone and there are very few data in total.
However, sufficient data exist to show the same sharp difference between the maritime
and continental Antarctica. For the mosses, growth rates decline from 10 mm year
1
in the
northern maritime to around 1 or <1 mm year
1
at the continental margin to an estimated
<0:2 mm year
1
in the Dry Valleys. A similar trend exists for lichens. Growth rates can be
measured relatively easily in the maritime (Sancho and Pintado 2004) and reach rates that
are similar to the largest known from anywhere in the world (Table 13.5, Figure 13.20).
However, growth is scarcely detectable in continental Antarctica. Figure 13.20 shows two
photographs of B. frigida taken 23 years apart in the Asgaard Range, Dry Valleys; growth is
detectable but is less than 0:01 mm year
1
. An intermediate rate is found for B. frigida of
0:06 mm year
1
over 42 years at Cape Hallett (Brabyn et al. 2005) and it appears that a
large cline in lichen growth rate exists from the northern Peninsula to central Victoria Land.
While there are photographic records showing colonization of man-made substrates by
lichens in the maritime, no such photographs exist for the continent (Kappen 1993a).
The longest available photographic record for analysis of change seems to be the map
produced by Rudolph in 1962 at Cape Hallett. This site has been remapped in 2003 and
substantial changes found in the cover of the three main vegetation types, algae, lichens, and
mosses (Brabyn et al. 2006). The largest change is an increase in the abundance of algae
indicating an improvement in water availability at the site. A series of long-term monitoring
sites have been established in the Ross Sea region to allow future monitoring of change
(Cannone 2006).
TABLE 13.6
Estimates of Growth Rates for Lichens and Mosses from Various Locations in Antarctica
Species Location
Growth Rate (mm y
1
unless Otherwise Stated) Comments
Lichens
Crustose spp. King George Island
(on whale bones)
40 mm in 78 year Kappen 1993a
Caloplaca sublobulata Livingston Island 0.86 Sancho and Pintado 2004
Acarospora macrocyclos Signy Island=
Livingston Island
1–3=0.72 Lindsay 1973=Sancho and
Pintado 2004
Xanthoria elegans Signy Island 0.2–0.5 Lindsay 1973
Rhizocarpon geographicum Signy Island=
Livingston Island
0.34=0.5 Lindsay 1973=Sancho and
Pintado 2004
Buellia latemarginata Livingston Island 0.87 Sancho and Pintado 2004
Buellia frigida Cape Hallett 0.06 Brabyn et al. 2005
B. frigida Asgaard Range
(Dry Valleys)
<0.01 Green unpublished
Mosses
Polytrichum alpestre Signy Island 2–5 Longton 1970
Calliergidium austrostramineum Signy Island 10–32 Collins 1973
Drepanocladus uncinatus Signy Island 11–16 Collins 1973
Bryum inconnexum Syowa Coast <1 Matsuda 1968
Bryum algens Mawson 1.2 Seppelt and Ashton 1978
Pottia heimii Taylor Valley <0.2 Green unpublished
Note: Sources of the data, which are representative, are given in the Comments column.
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Plant Life in Antarctica 421
ANTARCTIC PLANTS AS INDICATORS OF GLOBAL CLIMATE CHANGE
S
ITUATION
There has been increasing interest recently into the possibility of plants in Antarctica providing
early indications of global climate change, especially for temperature where any increase
would act to relieve both the geographical isolation and climatic constraints (Kennedy 1995,
Robinson et al. 2003, Hennion et al. 2006). This interest has been driven by the accepted
increases in temperature in Antarctica.
Global temperature rise is suggested to be greater and more rapid in polar regions. A
global increase of 0:03
C year
1
(0:2–0:5
C year
1
range) is predicted to be 0.5–0.7 times
larger in Antarctica. Temperature data from the past 45 years also show that a rise has
occurred; 0:067
C year
1
at Marguarite Bay (678 30
0
S), 0:056
C year
1
at Faraday Station
(658 S), 0:038
C year
1
at South Shetland Islands (628 30
0
S), 0:022
C year
1
at Signy Island
(638 43
0
S), and 0:032
C year
1
at Halley Station (768 S). As predicted by global change
models, the rise has been particularly large in winter temperatures. Although there is large
interannual variability the trend seems to be confirmed by glacial and ice shelf retreat. In the
remainder of Antarctica, particularly in the Ross Sea region, there is little evidence of any
increase in temperature (Doran et al. 2002).
Changes in vegetation have also been reported from the western Antarctic Peninsula.
Improved colonization by bryophytes and lichens has been seen, particularly in areas of newly
exposed ground however, the most impressive evidence comes from the expansion in abun-
dance of the two phanerogams (Fowbert and Smith 1994, Smith 1994). A study over 27 years
FIGURE 13.20 Photographs of lichens taken several years apart in the maritime and continental
Antarctic. Upper panels, Buellia latemarginata at Livingston Island (628 39
0
S) photographed in January
1991 (a), and January 2002 (b) (from Sancho and Pintado 2004); Lower panels, Buellia frigida at Mt
Falconer, Taylor Valley (778 35
0
S) photographed in December 1980 (c), and January 2003 (d). The scale
is about 1 cm.
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422 Functional Plant Ecology
has found large increases in populations of C. quitensis and D. antarctica both in Signy Island
(638 43
0
S) and the Argentine Islands (658 15
0
S). At Skua Island, Argentine Islands, as an
example, the population of D. antarctica was 190 plants in 1964 and 4868 plants in 1990
(Smith 1994). It is also clear that the rise in summer temperatures has led to increased
reproduction, germination, and survival.
PROBLEMS
At present the approach to the expected effects of global climate change appears to be fairly
simplistic and many papers simply demonstrate the effect of some factor on a lichen or moss
and then suggest how the species or community might change with altered climate. Change
might not be so simple to either predict or detect because several complexities are now known
to exist that compromise the simple approaches.
First, it is important that we know that change can actually be detected. Growth of
mosses and lichens is rapid in the maritime Antarctic but is much slower in the continent.
Recent studies show that change can be detected in both individual plants and communities
(Brabyn et al. 2005, Clarke et al. 2006, Brabyn et al. 2006) but that decades are required, not
single years. This is a major constraint on modeling change.
Biodiversity patterns in Antarctica are also more complex than originally thought. In the
Peninsula it is now confirmed (Peat et al. 2007) that a strong cline exists and that there is a
steady loss of species with increase in latitude. Such a situation is amenable in predicting the
effects of climate change, such as temperature rise, and the proof is in the already detected
changes in the phanerogams. The situation in the main continent is very different. The analysis
of lichen populations in the Ross Sea region suggests that the low precipitation has resulted in a
series of local microsites available for colonization and that this has been achieved by a sample
from the potential species. This is a more complex situation because the controlling factor
appears to be water availability and this is much more difficult to model than temperature.
An underlying assumption in any predictions of vegetation change with climate is that the
vegetation is in equilibrium with the local climate. At present there is little evidence to support
this assumption and some worrying indicators that it might not apply at least in some areas.
Work on collembolids has clearly demonstrated that population structure is determined by
past events rather than present conditions; for instance the formation of Ross Island (Stevens
and Hogg 2003), the presence of Lake Washburn in Taylor Valley 6000 years ago (Nolan et al.
2006), and the presence of relic populations that precede present glaciations and have
Gondwana affiliations (Stevens et al. 2006). The high endemic rate of lichens suggest an
ancient rather than recent origin (Peat et al. 2006), whereas the presence of species known
only from the Peninsula at Mt Kyffin, 848 S, Ross Sea, suggest relic populations in the same
areas as the collembolids (Tuerk et al. 2004). The somewhat enigmatic moss, B. subrotundifolium
(Seppelt and Green 1998) also has a Gondwana distribution known from the Ross Sea, New
Zealand, and Australia, all land masses that were earlier in contact. Soil communities in the
Taylor Valley are driven by legacy carbon also deposited at the time of Lake Washburn
(Lyons et al. 2000).
To the above uncertainties must also be added our lack of knowledge about dispersal
and colonization. Assuming that dispersal is possible (Marshall 1996) then the most likely
place to show the effects of warming is the Antarctic Peninsula, where a biodiversity cline
and warming are present. The second possibility is the ice-free areas that fringe the coast of
the continent at about the Antarctic polar circle. These are not warming but are so close to
having months with positive mean temperatures that community changes do not seem
unlikely in the future.
At present our best monitor for climate change seems to be the growth of lichens. This
shows two orders of magnitude change from 778 Sto628 S, a range matched by no other
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Plant Life in Antarctica 423
parameter (Table 13.6). This wide range almost certainly reflects the fact that the lichens and
mosses are not performing at their physiological optima under present ambient conditions so
that any increase in temperature would produce a concomitant rise in productivity.
There is growing evidence that a considerable propagule bank exists in Antarctic soils,
especially in the maritime zone. Use of cloches that can both increase the temperature and
water relations has resulted in substantial plant growth where none could previously occur
(Smith 1993, Convey and Smith 2006). Thus any area influenced by an improvement in
temperature or related water supply, especially in the Peninsula, will almost inevitably
become rapidly vegetated.
SUMMARY
It might appear, after an initial look at the literature, that there is very little evidence of
special adaptations by the lichens and mosses to Antarctic conditions. Lack of special
adaptation means that Antarctic species show much the same abilities as can be found in
temperate species. In addition, it appears that lichens in Antarctica are forced into the same,
but much rarer, microhabitat as those in temperate areas (Section ‘‘Active versus Inactive’’),
again suggesting little adaptation. This point has been made by previous authors
(e.g., Longton 1988a, Kappen 1993a). However, the rapid decline in species numbers from
the maritime to the continental zone and even more at higher latitudes indicates that, even if
no new adaptations are occurring, there is at least strong selection for a group of species that
can tolerate Antarctic conditions.
Ability to carry out photosynthesis at low and subzero temperatures and the related
tolerance to low water potentials seems to limit the major groups of plants (Schroeter
et al. 1994). Liverworts and cyanobacterial lichens seem to be excluded because they lack
this ability. It is particularly startling that cyanobacterial lichens are excluded because
cyanobacteria are extremely common in Antarctica producing substantial communities as
far south as the Scott Glacier, 848 S (Vincent 1988). However, the cyanobacteria are confined
to habitats where liquid water occurs, particularly the sides of ponds and streams. These
habitats cannot be occupied by lichens so the Antarctic is an unusual example of a situation
where the formation of a symbiosis (the lichen) does not extend the distribution of the
photobiont.
The large fall in productivity and growth rates between the maritime and the continent in
line with the decrease in precipitation, however, does indicate that the lichens and mosses are
predominantly controlled by the availability of water. The Antarctic is a cold desert with
water availability controlled not just by location but also by remaining frozen for long
periods. Plant growth is confined to those locations where precipitation is concentrated and
made available by warm conditions. This distribution control by water rather than by
temperature has been well explained by Kennedy (1993b). In essence, plants grow wherever
liquid water is reliably present as a result of glacier melt stream, by snow melt from stable
snow banks and by trapping of snow (e.g., by warm rock surfaces or fruticose lichens). It is
important that water, warmth, and light all coincide for plant growth. Kennedy (1993b)
makes the point that the Dry Valleys are a relatively warm area but no mosses or lichens (even
cryptoendoliths) can grow in them except where water is consistently present (Green et al.
1992). Antarctica is the coldest, driest, highest and windiest continent; the lichens and mosses
grow where it is warm, wet, low and protected.
Overall productivity is strongly influenced by the length of period when water is available
and the plants become, therefore, increasingly confined to areas of exceptionally good
microclimate. It is this strong link between microclimate, water availability and productivity=
growth that makes the system so potentially useful for monitoring global climate change,
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424 Functional Plant Ecology
especially temperature increase. The large change in biodiversity along the Antarctic Peninsula
covers the occurrence of months with mean temperatures above freezing. Even a small
increase in temperature will markedly alter the areas over which such months occur and
bring with it a marked community shift (Kennedy 1995).
It is becoming clear that there are two Antarcticas, the Peninsula and the main continent.
These differ in the controls on biodiversity distribution, there is a probably water unlimited
but temperature-determined biodiversity cline in the Peninsula compared to a, water-
controlled, temperature-independent, fragmented vegetation in the continent. The reverse
diel pattern of activity with the presence of very high light stress also seems to be confined to
the continent. Superimposed on this, and of particular worry to predicting climate change,
is the probable occurrence of population distributions on the continent that reflect past
events rather than present climate.
Antarctica will, therefore, remain in the future an important natural laboratory for
studying ecosystems under stress and the ecophysiology of cryptogams.
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