Introduction

Antarctica is a continent of superlatives and of contrasts. It is, for example, the coldest
and the driest and the windiest and the highest continent on earth. Yet it still supports a
range of organisms that extends from microbes and plants to animals. How the extreme
physical conditions evolved and how organisms are adapted to cope with them provides
an abundance of resource material for the science prescription. To this may be added
some unique aspects of Antarctic history and various socio-political elements which
reflect on broader issues, thus integrating, in Medawars terms, the art of the soluble
with the art of the possible.
As an educational resource, Antarctica is extremely broad in scope with the potential to
contribute to a number of study areas. The focus of this webpage and thus the majority
of the examples used, however, will centre on Antarctica as a resource for biology and
in particular on the various adaptations organisms living in Antarctica have evolved to
meet the unique conditions.
Map of Antarctica
Much can be made of these unique aspects of Antarctica. Who can fail but be aroused
by the extraordinary physical endeavours of Douglas Mawson, for example, when he
writes, having already buried his two sledging companions:
"On taking off the third and inner pair of socks the sight of my feet gave me quite a
shock, for the thickened skin of the soles had separated in each case as a complete
layer, and abundant watery fluid had escaped saturating the sock.........(T)here was
nothing to be done but make the best of it.........and with bandages (I) bound the old skin
casts back in place......Then on I went, treading rather like a cat on wet ground
endeavouring to save my feet from pain." (Mawson, 1915).
It might be added, as a casual aside, that this was on 11 January 1913. Mawson was
finally to walk into his base camp nearly four weeks later, having suffered and yet
survived some of the most extreme conditions on this planet.
Aims
The prime aim of this web page is to provide a focus for teachers interested in using
Antarctica as an educational resource. It represents an assemblage of primary and
secondary source material from both the written and the electronic media, both of which
should be accessed for further information.
Resource Potential
As an educational resource, Antarctica is extremely broad in scope with the potential to
contribute to a number of study areas (Table 1). Each of these different study areas will
be used as an example of how resource material relating to Antarctica can be applied to
advantage across different aspects of the educational spectrum. The focus of the web
page and thus the majority of examples, however, will centre on Antarctica as a
resource for biology.
Antarctic History
Reference to historical events associated with Antarctica offers a novel, interesting and
exciting platform for specific issues ranging from the extreme nature of the physical
environment (climate or geology, for example) to the need for environmental protection
(pollution or whaling, for example). A substantial summary of early Antarctic history is
provided by Harrowfield (1990).
Early History
Ancient Greeks named southern continent Anti-Arkitos.
Referred to later as Terra Australis Incognita (the unknown southern land)
The Exploratory Age (1772-1894)
James Cook (1772-1775) in the Resolution and the Adventure sailed to 71 10S
and aroused the interests of sealers and whalers on his return.
Thadeus von Bellinghausen (1819-21) in the Vostok and the Mirnyi reached 69
53 and sighted the Antarctic continent for the first time.
Dumont dUrville (1837-1840) in the Astrolabe and the Zelee reached 66 S.
Charles Wilkes (1838-1842) led a small flotilla of the Vincennes and 5 other
ships.
James Clark Ross (1840-1843) led the first British Antarctic Expedition in the
ships Erebus and Terror. They reached 78 17S and discovered the
Transantarctic Mountains, the Ross Sea, the Ross Ice Shelf, Victoria Land, Ross
Island, and Mts. Erebus and Terror.
The Heroic Age (1895-1915)
Henryk Bull (1895) managed a whaling expedition south in the Antarctica and
landed on the continent. Carsten Borchgrevink claimed to be the first, although
this was disputed by a 17 year old, Alexander von Tunzelman, who claimed he
leapt out to hold the boat steady.
Adrien de Gerlache (1897-1889) sailed south in the Belgica and over-wintered in
1898 when they became trapped in the pack ice in the Bellinghausen Sea. This
expedition is also noteworthy because among the crew was the young
Amundsen making his first trip south.
Carsten Borchgrevink (1898-1900) returned to Antarctica in the Southern Cross.
They over-wintered through 1899 at Cape Adare. The zoologist Nicolai Hansen
died on 14 October 1899 becoming the first person to be buried on the Antarctic
continent. During their stay they sledged to 78 50S, setting a new southerly
record.
Robert Falcon Scott (1901-1904) made his first of two trips south in the
Discovery. The ships was frozen in at Winter Quarters Bay (Ross Island) in 1902
and they built a hut nearby (adjacent to the current McMurdo Station). Able
Seaman George Vince lost his life on 11 March 1902 when returning across the
island with others from Cape Crozier. Scott, Edward Wilson and Ernest
Shackleton reached 82 17S on 30 Dec 1903. A party led by Armitage reached
the polar plateau for the first time and on their return discovered the Taylor
Valley, one of the Dry Valleys. The relief ship the Morning arrived in January
1903 under Captain Colbeck and left with Shackleton (who suffered from scurvy
on the push south) and a number of others. The rest stayed behind to overwinter
another season. In January 1904 the Morning returned with the Terra Nova
under Captain Mackay, the Discovery was freed using explosives, and the ships
left for New Zealand.




The cross erected on Hut Point (Ross
Island) in memory of George Vince.






Ernest Shackleton (1908-1909) returned in the Nimrod which was towed to the
pack ice by the Koonya to save coal. His group built a hut at Cape Royds on
Ross island. A party of six led by T. W. Edgeworth David climbed Mt. Erebus in
March 1908. One (Brocklehurst) had severely frost-bitten feet and had to be left
behind at an overnight staging point. The round trip from Cape Royds took 6
days. Shackleton, Jameson Adams, Eric Marshall and Frank Wild got to 88 23S
(148 km from the pole) on 9 January 1909, discovering the route up the
Beardmore Glacier to the Polar Plateau later used by Scott. Edgeworth David,
Douglas Mawson and Forbes Mackay reached the South Magnetic Pole on 16
January 1909.
Roald Amundsen (1910-1911) sailed south in the Fram. He ascended the Axel
Heiberg glacier with Hanssen, Wisting, Hassel and Bjaaland and reached the
South Pole (90 S) on 14 December 1911.

A Scott Base husky.

Robert Scott (1910-1913) returned in the Terra Nova and built their expedition
hut at Cape Evans. Edward Wilson, Henry Bowers and Apsley Cherry-Garrard
undertook "the worst journey in the world" to Cape Crozier in the middle of
winter. Scott, Wilson, Bowers, Lawrence Oates and Edgar Evans reached the
South Pole on 17 January 1912 (about a month after Amundsen) only to perish
on the return journey. Victor Campbell and other members of the northern party
were forced to overwinter in an ice cave on Inexpressible Island in Terra Nova
Bay. The Terra Nova returned on 18 January 1913.



Scott's hut at Cape Evans.


Douglas Mawson (1911-1914) led the Australian Antarctic Expedition sailing
south in the Aurora. This expedition is memorable for Mawsons epic sledging
journey in which his companions Belgrade Ninnis and Xavier Mertz lost their
lives.
Ernest Shackleton (1915-1917), having been beaten to the pole, planned a third
expedition (the Imperial Trans-Antarctic Expedition) using two ships. The Aurora
carried a group to lay depots to the bottom of the Beardmore Glacier, while the
Endurance was to deposit the crossing party on the other side of the continent.
However, the Endurance became trapped in the ice and sank in November 1915
and the crew of 28 took to the boats reaching the comparative safety of Elephant
Island. From here, Shackleton and five others sailed the James Caird 1300 km to
South Georgia and ultimately organised the rescue of the rest of the crew.
Meanwhile, the Aurora was taken out to sea in a blizzard and obliged to return to
New Zealand leaving a party of 10 under A. L. A. Mackintosh stranded on Ross
Island. This group laid the planned depots the following austral summer, but on
the return journey the Reverend A. P. Spencer-Smith died. Two others
(Mackintosh and Hayward) also perished when they attempted to cross the sea
ice from Hut Point to Cape Evans. The survivors were ultimately rescued by the
return of the Aurora with Shackleton on board.
The Mechanical Age (1916-1954)
The Ross Dependency was established and administered by the Governor
General of New Zealand on 30 July 1923.
Richard Evelyn Byrd (1929-1930 and 1933-1935). Sailed to Antarctica in the City
of New York and established Little America in the Bay of Whales.

In October 1956 an R-4D Dakota (Que Sera Sera) landed at the South Pole and
the construction of a permanent base began.

The International Geophysical Year (1957-58) increased scientific activities in
Antarctica with over 5000 personnel being involved. The year paved the way for
the Antarctic Treaty.

New Zealand established their Scott Base at Pram Point (3 km from McMurdo
Station) in January 1957.
Sir Vivian Fuchs, with Sir Edmund Hillary, led the Commonwealth Trans-Antarctic
Expedition. Hillarys team became the first since Scott to reach the South Pole,
while Fuchs team was the first to complete a trans-Antarctic crossing.
The Scientific Committee on Antarctic Research (SCAR) was established in
1958.
The Physical Environment
Geology
The evolution of organisms is closely linked to the evolution of their environment. The
major historical event which influenced the evolution of the Antarctic environment was
the splitting up of the ancient supercontinent of Gondwana (named after an ancient
Indian tribe known as the Gonds in conjunction with wana, meaning land).
The oldest rocks are Precambrian (about 3,000 million years old) and similar to those in
Australia, India, Africa and South America to which it was once joined. The
Transantarctic Mountains were laid down on the sea floor some 740 million years ago
(Precambrian) and subsequently uplifted and folded. By early Devonian times (350
million years ago) the early upfolded rock had been eroded to allow the sea to return
and the subsequent deposition of sediments (now fossiliferous) occurred. Global cooling
occurred during the late Carboniferous (300 million years ago) and early Permian times
(240 million years ago) and the future Antarctica was under glaciers. The climate
subsequently warmed and plant life became abundant in the later Permian forming what
is now recognisable as coal seams. Volcanic activity increased during the Triassic (190
million years ago) and during the early Cretaceous (178 million years ago) Gondwana
began to split up.
During the early Tertiary (45 million years ago) the Transantarctic Mountains began to
uplift, the Ross Sea was formed, and increased vulcanism saw the deposition of many
local landmarks. The development of the Transantarctic mountains led to the formation
of the Dry Valleys with the disappearance of their glaciers. Subsequent developments
have seen the shaping of the current landscape and the formation of soils from glacial-
deposited till in ice-free areas. Salts (derived from weathering and precipitation)
accumulate in the soils because there is little water to wash them away. Where moisture
exists, the soils can support microorganisms, mosses, lichens, fungi and a few
invertebrates such as springtails (Collembola).
The Splitting of Gondwana
The solid surface of the earth consists of a series of tectonic plates which can move
relative to each other. These plates are added to by basaltic material from the mantle of
the earth being extruded in mid-ocean ridges so that confluent plates are pushed into
collision at their margins. The collision sites are known as subduction zones since one
plate sinks under the other to return plate volcanic material back to the mantle. In early
geological times, collision of tectonic plates led to the crumpling up of the ocean floor,
examples of which can now be seen as folded mountain chains such as the Himalaya.
Movement of the tectonic plates over geological time (continental drift) has resulted in
the continuous restructuring of the earths surface. Part of the drift process resulted in
the splitting up of the ancient Gondwana supercontinent. The process began about 178
million years ago (early Jurassic) with the splitting of Antarctica from Africa, and ended
about 96 million years ago (late Cretaceous) with the separation of Australia and New
Zealand from Antarctica. Temperatures began to fall significantly around 35 million
years ago as the continent drifted away from South America and Australasia. Bradshaw
(1990) provides an excellent reference source for the early geologic history of
Antarctica.
Question
What is the evidence for the existence of Gondwana and how do we know it
separated to yield many of the major continents?
Physics and Chemistry
The Ozone Hole
Ozone is a molecular form of oxygen containing 3 atoms (O
3
) instead of the normal two
(O
2
). It is formed when an oxygen atom (the product of the breakdown of molecular
oxygen by sunlight) combines with molecular oxygen:
(i) O
2
+ hv O + O
(ii) O +O
2
O
3

Although it is only a minor component of the atmosphere (0.0001%), ozone plays an
important role in the upper atmosphere (the stratosphere, 15-50 km above the earth)
protecting us from the damaging effects of ultraviolet radiation, particularly UV-B (280-
320 nm). The amount of ozone in the atmosphere (recorded in Dobson units) can be
measured by spectrophotometry. The ozone hole is defined as an area of the
atmosphere where the ozone level is below 220 Dobson units. Prior to development of
the ozone hole the atmosphere above Antarctica is around 275 Dobson units. The
record low is 88 Dobson units recorded over the South Pole in 1993.
Damage to the ozone layer of the atmosphere was first reported by British scientists in
1985. They found that ozone over the Antarctic was significantly depleted during
October resulting in what is known as the ozone hole. On average, the hole grew in size
each year it was recorded until about 1994, when the increase began to level off. The
ozone hole now spreads over about 26 x 10
6
km
2
, which is roughly the area of North
America.
The increase in size of the ozone hole is due to the presence of synthetic chlorine-
containing gases which are transported high into the stratosphere (especially over the
South Pole in spring) where they are eventually broken down to yield a chlorine atom
that promotes the destruction of ozone. Bromine also contributes to the breakdown of
ozone.
(iii) O
3
+ Cl O
2
+ ClO
When a free atom of oxygen collides with the chlorine monoxide, molecular oxygen is
produced releasing the chlorine atom to attack more ozone:
(iv) O + ClO O
2
+ Cl
In this way, one atom of chlorine can destroy about 100,000 molecules of ozone. This
has led to the perturbation of the natural balance of ozone in the atmosphere.
The ozone hole disappears in the austral summer because the polar vortex which
creates the spiralling of air over the South Pole in spring breaks down allowing ozone-
rich air from other latitudes to mix with that in the polar region. The air within the polar
vortex is very cold (about 80 C) and this allows polar stratospheric clouds to develop
which are also crucial to ozone depletion since on their surfaces active chlorine is able
to form from intermediate reservoir molecules. Obviously the chemistry of ozone
depletion is far more complex than is summarised here.
Typical among the synthetic chlorine-containing gases are the chlorohydrofluorocarbons
(CFCs) such as freon, which is used in refrigerators and air conditioning units. Because
they are relatively stable compounds, the CFCs have been accumulating in the
environment since they were first introduced. Even if all CFC and related chemical
production was to cease immediately, it would still take until about 2050 AD before the
ozone hole disappeared.
Excessive depletion of ozone in the atmosphere results in more ultraviolet radiation
reaching the earths surface. High energy UV-B radiation is particularly damaging,
capable of altering DNA leading to cancers such as malignant melanoma. UV-B can
also damage proteins, and in plants it is capable of bleaching chlorophyll such that it no
longer functions as an energy absorbing molecule for photosynthesis. Because UV-B
can penetrate clear sea water to a depth of about 20 m it can also impact on the
productivity of phytoplankton which are at the base of the marine food web.
It is thought that the ozone hole will remain roughly similar in size in the near future with
a possible decrease in the next 30-50 years as chlorine levels in the atmosphere
decline.
Aurora Australis
The aurora australis, or southern lights, arises from plasma (ionised gases) in the solar
wind (a stream of plasma flowing out from the sun) penetrating the earths atmosphere
and colliding with nitrogen and oxygen to emit green and red light respectively. The ions
of the solar wind are concentrated at the polar regions along the earths magnetic field
lines and thus auroras are seen predominantly in these localities. An aurora can
generate up to 10
13
watts of electricity providing enough light to read by.
Geography
Antarctica as a Wilderness Region
The Antarctic continent including all the islands and ice shelves covers approximately
13,661,000 km
2
in area. This is about 50x the area of New Zealand. The continental
coastline measures about 17,968 km, and the Antarctic continental shelf covers about
4x 10
6
km
2
at a mean depth of 350 m (nearly twice that of other shelves). Antarctica is
the highest continent in the world with an average height above sea level of 2500 m.
The highest peak is Vinson Massif (4897 m). Antarctica is almost entirely covered by
ice, with exposed rock accounting for only about 0.4% (48,310 km
2
) of the continent.
The ice cap has an average thickness of about 2450 m, reaching to 4,776 m at its
maximum. There are two active volcanoes in Antarctica, one on Deception Island (off
the Antarctic Peninsula) and the other (Mt. Erebus) on Ross Island, and around 70
extinct ones. Mt. Erebus (3794 m) contains a permanent molten lava lake in the summit
crater. The solidified basaltic lava of Mt. Erebus is known as kenyte.
The Dry Valleys
The region known as the Dry Valleys is only one of a number of lesser known "Antarctic
deserts". It encompasses the Taylor, Wright and Victoria Valleys and occupies an area
of about 2500 km
2
. Strong, dry winds (relative humidity <10%) sweep down the valleys
from the polar plateau serving to keep the rocks bare of snow. Microorganisms, fungi,
plants and invertebrates can only survive in the Dry Valleys where they can find
moisture. An interesting adaptation involves organisms such as certain cyanobacteria,
algae and lichens which can actually live and grow inside sandstone rocks of the Dry
Valleys, where they form what are known as "endoliths". Numerous mummified seals
(mostly crabeaters) and some mummified Adelie penguins are also found in the Dry
Valleys. It is thought that these carcasses are of animals that became disoriented and
headed inland instead of seaward, eventually to die of dehydration up to 40 km or so
from the coast. The carcasses are freeze-dried in excellent condition by the climate in
the Dry Valleys, but they eventually erode as the wind takes its toll.


The Wright Valley.



Question
The Dry Valleys are technically deserts. In what ways do they differ from more
typical deserts such as the Sahara?
The Ice Sheets
The Antarctic ice sheet (or ice cap) is composed of the East (or Greater) and West (or
Lesser) Antarctic ice sheets which effectively merge into one. The eastern sheet is
larger, thicker and terrestrial, since it rests on a land base. The Transantarctic
Mountains are on its western side (with reference to the Greenwich meridian). The
western sheet is smaller, thinner and marine since its base is on the ocean floor. The
Antarctic ice sheet covers about 99% of the continent extending over 14 x 10
6
km
2
to a
maximum depth of 4700 m. It contains about 30 X 10
6
km
3
of ice and results from the
continuous accumulation of snow which is packed down under its own weight to form
dense glacial ice.


An ice cave which falls away to a crevasse
on the right hand side.


The glacial ice sheet flows in ice streams under gravity towards the sea (at up to 1 km
per year) where it forms ice shelves that float on the sea while retaining a connection
with the ice sheet. Since the ice shelves float they are able to flow faster, at up to 3 km
per year. The Ross Ice Shelf covers 490 x 10
3
km
2
(nearly twice the size of New
Zealand) and the Filchner-Ronne Ice Shelf about 449 x 10
3
km
2
.
At their margins the ice shelves break into tabular icebergs which drift off and eventually
melt adding their fresh water to the sea. About 1450 km
3
of icebergs are calved off the
ice shelves each year and drift in the circumpolar currents at about 13 km per day. The
largest iceberg ever recorded measured approximately 335 x 97 km which covers an
area about half the size of Tasmania. Since about 80% of all of the worlds freshwater is
contained within the Antarctic ice, icebergs represent a potential source of freshwater
for the desert regions of the world. Although average-sized icebergs can last for 2 or
more years (depending on their volume, shape and location) they are seldom found
north of 55 S, however, because of the temperate waters.


A tabular iceberg adrift in McMurdo Sound.



There is increasing concern that global (or local) climate changes resulting from the
accumulation of greenhouse gases (e.g. carbon dioxide; nitrous oxide) could promote
the melting of the ice shelves putting many countries at risk through rising sea water
levels. Temperatures on the Antarctic Peninsula, for example, have risen 2.5 C since
the 1940s. Over the last 100 years the global sea level has risen by between 10 and 25
cm. Although an unlikely event, if the entire Antarctic ice sheet were to melt, the sea
level would rise by about 60 m.

The Ross Ice Shelf
The Ross Ice Shelf varies in thickness from over 1000 m where it is fed by glacial ice
from the ice sheet to less than 100 m at the ice front. On its seaward side it calves
icebergs at a rate of about 150 km
3
per year, and it also loses volume by bottom melting
at a rate of about 2.5 m per year. One of the largest icebergs recorded from the Ross
Ice shelf in modern times appeared in the spring of 1987 and measured about 154 km
long x 35 km wide.


A grounded iceberg off Ross Island.



Sea Ice
The sea around the ice shelves is often frozen to form 2m thick sea ice which can
extend from 4 x 10
6
km
2
in late summer (February) to 22 x 10
6
km
2
before the thaw
begins (September). The sea ice forms an insulating blanket over the sea and restricts
light penetration which slows the growth of marine phytoplankton.

Drilling through the sea ice (over 1 m
thick)
in Winter Quarters Bay where Scott
anchored the Discovery when
overwintering on his first expedition.


The formation of sea ice is an annual phenomenon although some parts do not melt
during the following summer. It begins with the freezing of the sea surface and the
formation of "frazil ice" which later becomes slushy "grease ice". The patches of grease
ice consolidate into loosely packed, flat, round plates known as "pancake ice". These
thicken and coalesce to form larger slabs or floes which merge to form a solid sheet
("pack ice").
Frazil ice can also form in the water column giving rise to ice crystals and larger
platelets (10-15 cm diameter) which float up and accumulate on the undersurface of the
sea ice where they form "platelet ice". In shallow water, ice can form on both the upper
and lower surfaces, the latter giving rise to what is known as "anchor ice".
Sea ice begins to form in the Ross Sea in March/April and is usually dispersed by mid
summer to leave open water again, except where pockets of fast ice remain in bays and
inlets. Pockets of open water (polynas) persist throughout the winter in a number of
locations.
Question
What issues does the presence of sea ice raise for the marine ecosystem in
Antarctica?
Sub-glacial Lakes
A huge lake covering some 14,000 km
2
land reaching 670 m in depth lies underneath
the East Antarctic ice sheet in the vicinity of the Vostok research station. This lake,
known as Lake Vostok, is one of 70 sub-glacial lakes on the continent. They are thought
to form when the temperature beneath the ice sheet reaches melting point (about -1.5
C), probably as a result of the friction generated as the ice moves over the land mass
under immense pressure. Because Lake Vostok is so large the ice sheet floats on it
presenting a flat surface that enabled it to be mapped from space by satellite.
An attempt to drill through the ice sheet began in 1989, but was stopped in 1998 about
120 m above the lake surface (at a depth of 3623 m) to allow scientists to consider the
consequences of breaching. Studies of the ice core showed it to be about 500,000
years old at this depth. Viable microorganisms have been isolated from parts of the ice
core estimated to be 200,000 years old raising the possibility that the lake and its
contents may provide information relevant to life in extreme environments such as that
which existed when life first evolved. Unfortunately, most of this information can only be
tapped by penetrating the lake itself. This presents significant problems since once the
lake is penetrated contamination of its waters is inevitable. Ice drilling, for example, is
enhanced by the use of a drilling fluid (the Russians typically use a mixture of aviation
fuel and freon) with significant potential to contaminate the lake. Clearly, considerable
thought is required before any strategy to penetrate the roof of the sub-glacial lake is
put in place.
Question
Should Lake Vostok be breached? What are the relative advantages and
disadvantages of doing so?
Climate
Temperature
The Russian base at Vostok (78.5 S, 106.9 E), which is 3488 m above sea level, is
the coldest place on earth with a mean annual temperature of around 55 C. The
coldest recorded temperature at Vostok was 91 C in 1997. This is far colder than dry
ice, which has a temperature around -70 C. The South Pole (90 S), which is at 2,835
m, has a mean annual temperature of about 49 C.
Why is Antarctica so Cold?
First, the sun is always relatively low in the sky over Antarctica so its surface
receives less energy. On Midsummer's day at the South Pole, for example, the
sun is only 23.5 above the horizon and a shaft of sunlight is spread over a much
larger area than an equivalent one would be on the equator.
Second, the sun does not rise at all in regions south of the Antarctic Circle for
part of the year. At the latitude of the Antarctic Circle there is one day a year of
total light (21 December) and one of total darkness (21 June), whereas at the
South Pole, there are 6 months of daylight and 6 months of darkness. On
average, Antarctica actually loses more energy by infrared cooling than it gains
from the sun.
Third, the whiteness of the ice sheets and the sea ice (which together can extend
over 35 x 10
6
km
2
in the winter) results in the reflection of up to 85% of the
incident polar radiation.
Fourth, Antarctica is the highest continent on earth (average height 2,500 m).
Away from the coast, the temperature decreases by about 1 C for every 100 m
rise in height.
Fifth, the Antarctic Circumpolar Current flows completely around Antarctica
largely preventing other warmer ocean currents from exerting significant effects.
Finally, Antarctica is an isolated continent without significant neighbouring land
masses which can have a warming effect on the climate.
Wind
Cape Denison, the "home of the blizzard" where Douglas Mawson built his hut in 1912
is probably the windiest place on earth. During one spectacular and persistent hurricane
Mawson recorded "a terrific series of Herculean gusts..... in the order of 200 mph" (320
km h
-1
). The highest recorded wind velocity is 327 km h
-1
which was measured at the
French base of Dumont dUrville in July 1972.
Why is Antarctica so Windy?
The powerful gusts which caused Mawson so much trouble are the result of katabatic
winds (from the Greek katabasis, going down) which arise when cold, dense air lying
less than a few hundred metres off the surface at the highest levels of the Antarctic ice
sheets flows down towards the coast under gravity. Near the coast these winds can
reach the tremendous speeds Mawson experienced, in some places averaging over 70
km h
-1
for the year. The onset of a katabatic wind is very rapid, it seems "to come from
nowhere" and can disappear equally as fast. A temperature increase is usually
associated with the arrival of a katabatic wind due to the sweeping away of the cold air
layer that forms under calm conditions.
Antarctic Biology
The Southern Ocean Ecosystem
The Southern Ocean, which is entirely surrounded by sea, is a mass of water and ice
covering approximately 36 x 10
6
km
2
which represents about 10% of the world's
oceans. The northern limit of the Southern Ocean is set by the Antarctic Polar Front
(Antarctic Convergence), identifiable as a 40 km wide ocean band where a 2-3 C
change in temperature of the sea surface occurs. This relatively abrupt change in water
temperature occurs where dense, cold polar water meets and flows under less dense
warmer water from the tropics. The Antarctic Polar Front separates the Southern Ocean
from the Atlantic, Pacific and Indian Oceans which border it to the north. Water in the
Southern Ocean moves eastwards (clockwise) around Antarctica carried by the
Antarctic Circumpolar Current, the largest ocean current in the world. A counter-current,
the Antarctic Coastal Current, flows westward closer to the continent.
The Southern Ocean Food Web
The Southern Ocean food web describes the flow of energy within the Southern Ocean
ecosystem from microbes to the top predators. Marine phytoplankton (the primary
producers) use energy from the sun to make simple organic molecules by
photosynthesis. The phytoplankton fix about 50 tonnes of carbon (from CO
2
) per 100
km
2
of sea surface each year. Because of their dependence on light, phytoplankton
growth is restricted to surface waters where there is little sea ice (which would otherwise
limit light penetration), and it is influenced by seasonal variations in sunshine. A small
amount (<10%) of the organic molecules produced by photosynthesis are recycled by
other microorganisms which may drop to the sea floor where they fuel bottom-dwelling
organisms. The major proportion passes through the food web at the hub of which is the
Antarctic krill (a key herbivore) and a variety of zooplankton. Predators of krill such as
the baleen whales, crab eater seals, penguins, fish and squid are the primary predators
of the food web. Higher order predators include the killer whale, leopard seal and birds
such as the wonderful skua.
Question
Sketch a summary diagram highlighting the major features of the Southern
Ocean food web. How does this relate to the effects of global environmental
pollutants?
Krill
The Antarctic krill, of which there are several species, is a shrimp-like crustacean
growing to about 5 cm in body length. Krill (the name is derived from a Norwegian
whaling term meaning "small fry") are widely distributed around the Antarctic continent.
Euphausia superba dominates north of the pack ice, while the smaller E.
crystallorophias is the principle krill species under the ice. E. superba form dense
swarms, typically hundreds of metres across and 20 or so meters deep. The largest
swarms can spread over several square kilometres and may extend to a depth of 200
metres. Swarms of such dimensions may contain up to 10 x 10
3
tonnes of krill which
makes them an extremely abundant food supply. It has been estimated that there are
over 6 x 10
11
individual krill with a biomass of around 250-600 x 10
6
tonnes but,
because of their patchy distribution, these figures may not be particularly reliable.
Although labelled as the dominant herbivore in the Southern Ocean food web, krill are
actually omnivores feeding on the zooplankton as well. The life history of E.
crystallorophias is not well known and further study of this species remains a great
challenge to scientists.



The Antarctic krill - Euphausia superba.
For more information about krill visit Uve Kils'
website from where this image is sourced.
Adaptations
Krill display a number of interesting adaptations, particularly behavioural ones such as
swarming and movement in the water column which may be related primarily to their
feeding habits. Perhaps their most profound adaptation relates to their survival during
winter when food supplies are relatively scarce. Instead of building up large fat
reserves, krill reduce their size capitalising on their own body proteins as a source of
energy. This reduction in size presents a special problem for animals which have
exoskeletons. The problem is overcome in krill by retaining the ability to moult into
adulthood. The ability to increase or decrease size depending upon food supply enables
the krill to survive during the winter. This is further enhanced by their omnivorous rather
then strictly herbivorous character thus maximising available food over the winter
months.
Fish
The modern Antarctic fish fauna is exclusively marine, with 203 bottom-dwelling species
and 75 mesopelagic species being recognised (reviewed in Eastman, 1993). This
represents about 1% of all known fish species. The fauna is highly endemic (about 88%
of the species are restricted to Antarctica) and of the bottom-dwelling species, 104
(51%) belong to a single suborder, the Notothenioidei. The fossil fish fauna shows that
Antarctic fish have undergone a considerable reduction in diversity since the early
Tertiary, but the reasons for this are not clear, particularly since there are no
notothenioids in the fossil record to date. The dominance of notothenioids presumably
reflects the presence at some early stage in their evolution of an ancestral species
which overcame the difficulties of survival at low temperatures. Subsequent speciation
and evolution of this ancestral species resulted in the current notothenioid fauna
(monophyletic radiation). The suborder Notothenioidei comprises six families: the
Nototheniidae (Antarctic cod), the Harpagiferidae (spiny plunderfishes), the
Bathydraconidae (dragonfishes), the Channichthyidae (icefishes), the Bovichthyidae
(thornfishes), and the Artedidraconidae (plunderfishes).


The nototheniid - Trematomus bernacchii.
Questions
After whom was this fish named?
How was he involved in early exploration of
the Antarctic?
Question
Explain the term "monophyletic radiation" with reference to the evolution of the
Antarctic fish fauna. What lines of evidence are available to support the concept
of monophyletic radiation?
Adaptations
During their evolution the Antarctic fishes have adjusted to cope with the Antarctic
environment (reviewed in Macdonald and Montgomery, 1990). None possess a gas-
filled swimbladder (used for buoyancy) which suggests that the ancestral species was
probably a bottom dweller. Some of the modern species such as Pleurogramma
antarcticum have become secondarily pelagic, decreasing their densities by
incorporating lipid into their tissues and developing a cartilaginous skeleton with hollow
vertebrae. Many notothenioids have pelagic larvae which can feed directly on plankton.
Since these are low in the food chain, the larvae benefit from a larger, potentially more
available food source.
Although obviously cold (-1.86 C at high latitudes to +5 C near the Antarctic Polar
Front), the waters of Antarctica are relatively stable (seasonal variation is typically <
0.2 C) and local fish species have evolved to become intolerant of temperature
changes. The concentration of salts in sea water depresses its freezing point to about
1.9 C, while that in fish blood depresses it to about 0.8 C. To survive at near freezing
temperatures fish contain various types of antifreeze in their blood, reaching a
concentration of about 35 mg/ml. Fish of the species Pagothenia borchgrevinki contain
8 glycopeptide antifreezes of similar structure of different molecular weight (2,600-
33,700 daltons). Each has a repeating tripeptide (3 amino acid) backbone (threonine-
alanine-alanine) with a disaccharide extending from the threonine residue. The
antifreeze glycopeptides act by binding to small ice crystals in the fish to prevent further
growth. Although present in most body fluids, there is no anti-freeze present in the
urine. This is a result of specialisation of the kidneys which do not contain glomeruli.
Filtration of the blood takes place by active secretion through the walls of the kidney
tubules, thus preventing loss of anti-freeze glycoproteins.


The Nototheniid - Pagothenia
borchgrevinki.
Questions
After whom is this fish named?
What was his contribution to Antarctic
exploration?
The viscosity of blood increases at lower temperatures, increasing the energy required
to pump it around the body. Many Antarctic fish are adapted to this situation by having
fewer red blood cells (the predominant blood cell type), thus effectively thinning the
blood. In temperate fish the corresponding reduction of red blood cell haemoglobin
would significantly lower the amount of oxygen carried to the tissues, but the increased
solubility of gases at low temperature allows Antarctic fish to carry more oxygen
dissolved directly in the blood. The solubility of oxygen in sea water at 0 C, for
example, is 0.83 vol. % per atmosphere of air, while its solubility at 20 C is 0.53 vol. %.
The channichthyids (icefish) are unique among vertebrates in that they have a deleted
gene for haemoglobin and thus they are totally dependent on the increased solubility of
gases at low temperatures to transport oxygen in their blood plasma. The oxygen
carrying capacity of icefish blood is about 0.7 vol. %, which is only 10 or so of the
amount carried by the blood of haemoglobin-containing Antarctic fish. Transport of
oxygen in icefish is facilitated by a number of anatomical and physiological adaptations
including a large heart (similar in weight to that of a small mammal) with a large stroke
volume (6-15 x that of other teleosts) increasing cardiac output at low heart rate and low
ventral aortic pressure. They also have a high blood volume (2-4 x that of other
teleosts), large diameter blood vessels to lower resistance, low blood viscosity, well
vascularised gills and a scaleless skin to maximise the extraction of oxygen from the
sea water, and a relatively low average metabolic rate minimising the demand for
oxygen. The icefish are so well adapted they appear not to be compromised by their
lack of haemoglobin and they can lead active, predatory lives.
Birds
There are a considerable number of avian species which visit Antarctica, but relatively
few breed on the continent. Of those that do, discussion will be restricted to the skuas
and the penguins.
Skuas
There are five currently recognised species of skua. The two which dominate on the
Antarctic continent are the South Polar skua (Catharacta maccormicki) and the brown
skua (C. lonnbergi). The South polar skua is widely distributed over the mainland,
whereas the brown skua is restricted to the Antarctic peninsula. Interbreeding is
possible where the distributions of the two species overlap.
Lieutenant Evans (who went south with Scott) wrote that their greatest value to the early
explorers was their tasty and nourishing flesh, reminding him somewhat of wild duck.
He was either possessed of a wonderful imagination or extraordinary digestive powers.




An adult south polar skua (Catharacta
maccormicki ) with chick.
The south polar skua.
The south polar or Antarctic skua is also known as McCormicks skua, after Dr
McCormick (the surgeon on Ross expedition), who history records as the first
person to shoot one. It is a handsome bird with a light-brown head and breast
progressing to a darker brown back and posterior. Its webbed feet are clawed
and it has a vicious looking curved beak. It is undoubtedly the most southern
existing animal species, one bird apparently following Amundsen almost to the
South Pole itself. Those nesting near penguin colonies feed mostly on
unattended penguin eggs and chicks when in season, but most feed at sea on
Pleurogramma antarcticum. Skuas nesting near research stations are known as
skilled scavengers. Young (1994) provides an excellent summary of the
relationship between penguin and skua, predator and prey.
The south polar skuas on Ross Island leave the region over winter, migrating to
Japan and the Alaskan coast, and return in late October. The first eggs are laid in
mid-November and the first chicks hatch about a month later, leaving the colony
as fledglings in late March. Two eggs are generally laid, and both usually hatch,
but within a day or two of hatching the second chick is attacked by the stronger
first chick and driven from the nest where it falls as prey to other adult skuas.
This display of siblicide is apparently not shown by subantarctic skuas, and may
be related to food availability. The skua is the scavenger of the south,
congregating around scientific bases seeking food scraps. They also haunt
penguin rookeries seeking eggs, but also attacking chicks. At sea, the skua feeds
on Antarctic silverfish Pleurogramma antarcticum.


Sibling south polar skua chicks.
The first born (and the larger of the
two) is harassing its younger sibling
which will be driven ultimately from
the nest. Once excluded it will fall
prey to other adult skuas which are
always on the lookout for an easy
meal.
Penguins
There are 17 species of penguins distributed over six genera, all of which are found in
the family Spheniscidae. They are all flightless, pelagic seabirds and vary in size from
the Little Penguin (Eudyptula minor) which weighs just over 1 kg and is around 40 cm
tall to the Emperor (Aptenodytes forsteri) which is a hefty 30 kg and reaches 115 cm in
height. Only two penguins are restricted to the Antarctic, the Adelie (Pygoscelis adeliae)
and the Emperor.
The Adelie penguin
The Adelie penguin (Pygoscelis adeliae) at about 5 kg is among the smaller of
the penguins. It is easily identified by its distinctive white-rimmed eyes set in a
black head. It was named by the French explorer Dumont dUrville after his wife.
It is the most numerous of the Antarctic penguins and breeds further south than
most, the most southerly colony of about 4000 birds being at Cape Royds on
Ross Island (77 30 S, about 1350 km from the South Pole). The largest colony
is at Cape Adare where there are about 400,000 birds.


Adult Adelie penguin ( Pygoscelis
adeliae ).
The Adelie is easily recognised by the
white ring around its eyes.
The first Adelies come ashore around the third week of October and egg laying
peaks in the middle of November. At the start of the breeding season there is still
a great deal of solid ice between the birds and their nesting sites and distances
of 20-40 km may have to be traversed over the ice mass. Egg incubation lasts
about 34 days, with most chicks thus appearing in mid-December. The adult
birds survive the early weeks of the season without feeding, the females through
to egg laying and the males through the first 15-20 days of incubation. Food for
the chicks, mostly krill, is provided after hatching by their parents which spend
large parts of the day hunting at sea. Adults typically travel 20-30 km a day in
search of krill swarms which they collect in short dives of 10-40 m depth,
occasionally extending to 170 m. Females have been tracked to over 340 km
from the colony, but distances around 10 km are more common when rearing
chicks. Within two weeks the chicks have put on enough weight to insulate
themselves against the weather and they can wander around the colony. The
first chicks are ready to go to sea in late January, having lost their coat of fluffy
down, and in late-February the adults begin to moult to get a new set of
protective feathers. Before moulting begins the adult Adelies return to the sea for
2-3 weeks.
The poor Adelie provided a source of blubber for fuelling the fires of the early
explorers and, along with its eggs, also made its way to the dinner table. The
indefatigable Lieutenant Evans wrote that the adults tasted "quite like hare, and
(are) much improved by red currant jelly". The Adelie was not to everyones
taste, however, and Frederick Cook (a member of de Gerlaches 1897
expedition) described it more like "a piece of beef, odiferous codfish and a
canvas-backed duck roasted in a pot with blood and cod liver oil for sauce".
Somehow or another this latter description seems more likely!

The emperor penguin
The Emperor penguin (Aptenodytes forsteri) is a big bird, over a metre tall and
weighing around 30 kg. They can dive deeper than any other penguin reaching
depths in excess of 450 m to find their preferred food of fish and squid. They also
range widely in search of food, often travelling 150-1000 km in a single foraging
trip. There are about 400,000 Emperor penguins breeding at a number sites on
the Antarctic continent. The breeding season of the emperor penguin is so long
that it has been advanced into the winter. At Cape Crozier, breeding birds
congregate in March/April, the eggs are laid May/June, and the chicks hatch in
July/August. Soon after laying her single, large egg (460 g), the female returns to
sea for the next 64 or so days, leaving the male to incubate the egg alone in the
most hostile environment imaginable. As well as having an extensive range of
physiological adaptations to cope with the extreme conditions, the Emperors
huddle together at about 10 birds per square meter to share body heat. The
Emperor does not make a nest on the bare ice, but instead incubates the egg
between a loose outer fold of abdominal skin and a patch of bare belly skin which
is highly vascularised.

An adult Emperor penguin.
The male does not eat from first arriving on the ice until it is later relieved from
incubating duties by the returning female. This covers a period of about 115 days
in which the male can lose up to half its weight. In order to return to the brooding
site, the female must walk back from the ice edge over as much as 100 km or
more since at this time of year the ice is almost at its maximum extent. When the
female reaches the tightly packed colony, it calls out to its partner to re-establish
their bond. The chick could have hatched while the female was at sea, in which
case the male has to feed it until the female arrives. The male can do this for up
to two weeks by regurgitating a rich secretion from its crop, but if the female is
delayed much beyond this time the male has to abandon the chick and begin its
own walk to the ice edge. After the female has found her partner it immediately
begins to feed the chick. It then takes over the incubating duties to allow the
bedraggled male to return to the sea. Over the next 5 months or so the parents
travel back and forth over the ice to forage and bring back food for the chick
which rapidly puts on weight. By the time the chicks are 5-8 weeks old they have
well and truly outgrown their brood pouches and they leave their parents to
gather in crches. At the end of this time the parents return to the sea in
preparation for moulting and they are soon followed by the abandoned chicks,
now about 21 weeks old.

In order to shed some light on the embryonic development of the Emperor, three
members of Scotts last expedition (Edward Wilson, Henry "Birdie" Bowers and
Apsley Cherry-Garrard) undertook "the worst journey in the world" travelling
about 110 km on foot from their hut at Cape Evans to an Emperor penguin
colony at Cape Crozier (Cherry-Garrard, 1922). They left Cape Evans on 27
June 1911 and made camp at Cape Crozier on July 15, manhauling 343 kg on
two sledges. Their journey took place in the permanent dark of an Antarctic
winter in temperatures which dropped to -61 C and in tremendous gales which
at one stage ripped their camp apart and blew away their tent. Without their tent,
the three explorers would almost certainly have died of exposure on their return
journey. By a stroke of extraordinary good luck they were able to find it in
undamaged condition when the gale subsided. The three ultimately made it back
to base camp at Cape Evans, although Wilson and Bowers were later to perish
with Scott on the ill-fated assault on the Pole.
Adaptations
Like all animals living in Antarctic, birds are adapted to cope with the extremes of the
environment. This is demonstrated clearly by the penguins which have a thick
thermoprotective layer of subdermal fat to maintain body heat while also streamlining
the body for swimming. Their short, overlapping feathers trap insulating air when diving
and promote insulation even under windy conditions. Heat loss through the legs and
feet is minimised by keeping them tucked up against the warmth of the body and by
having heat exchangers in which heat is transferred between intertwined arteries and
veins. Many penguins such as the emperors also huddle together in extreme conditions
to retain body heat. Their efficiency in heat retention is such that they have evolved
special adaptations for those occasional warm sunny days that do occur in Antarctica.
These adaptations take the form of blood vessels which come close to surface on the
almost featherless inside surface of their flippers. Thus in sunny conditions, or when
otherwise hot, penguins can radiate metabolic heat by raising their flippers to expose
the undersides.
Seals
The seals (Order Pinnipedia) are distributed over three families: the families Phocidae
(true seals) and Otariidae (the eared seals) are found in Antarctica, whereas members
of the family Odobenidae (the walruses) are not. There are six species of Antarctic
seals. These include the Antarctic fur seal (Arctocephalus gazella) which represents the
Family Otariidae, and five phocids represented by the elephant seal (Mirounga leonina),
the Weddell seal (Leptonychotes weddelli), the Ross seal (Ommatophoca rossi), the
leopard seal (Hydrurga leptonyx) and the crabeater seal (Lobodon carcinophagus).
Weddell seals
The Weddell seals (Leptonychotes weddelli) are an ice-breeding species which
reach about 3 m in length and which weigh up to 400-500 kg. There are about
800,000 Weddell seals in Antarctica where they live all year round on the fast ice.
They are the only mammal which over-winters in the Antarctic, humans excluded,
and they breed further South than any others. At White island, about 20 km from
the barrier edge, there is an apparently self sustaining population of Weddell
seals cut off from the seasonally open water of McMurdo Sound. Assuming that
Weddell seals can travel no more than 12 km under the water on one breath, and
given that there are apparently no breathing holes on the Ross Ice Shelf in this
area, the inevitable conclusion is that they must be an isolated population. DNA
testing is required, however, to resolve this possibility.


Weddell seals ( Leptonychotes
weddelli ) on the sea ice in
McMurdo Sound.
The Weddell seal feeds mostly on fish and squid, diving to depths greater than
700m to find them. Dives can last over 60 min, but most feeding dives are short
(<30 min) and shallow (200-400 m). The average meal size is a whopping 23 kg
(thats 257 quarter pounders!). A Weddell seal was once seen to catch a 1.5 m
long Dissostichus mawsoni estimated to weigh 31 kg which was brought to the
surface and consumed over three hours. The hunting territo
Adaptations
All the Antarctic seals display similar general adaptations to the local
environment, but each species also has unique adaptations reflecting their
occupancy of distinct ecological niches. Weddell seals feed primarily on fish and
have peg-like molar and pre-molar teeth to grip their prey. During the winter they
live under the ice (where it is warmer) and they thus need to keep their breathing
holes from freezing over. This is achieved by having their canine and second
incisor teeth modified for ice cutting which is undertaken by dragging the upper
teeth (which project forward) from side to side on the rim of the ice hole. This
action wears down the upper canines and second incisors, and tooth wear with
associated ulceration is probably an important factor contributing to the relatively
short life span of Weddell seals. Crabeater seals, found on the pack ice in the
north of the Ross Sea, have remarkable five-pointed molars which interlock when
their jaws are closed to form a strainer capable of sieving krill. The common
name "crabeater" is thus misleading as they eat krill and not crabs. Leopard
seals, also common on the pack ice, have interlocking, tricuspid molars ideally
suited for sieving krill, but they are also accomplished predators preying on
penguins, other seals (especially young crabeaters), fish and squid and have
long and slender canine teeth. Their lower jaw is relatively massive as is
common in generalised predators.
More generally, the Antarctic seals have a thick insulating layer of blubber which
is poorly vascularised, thereby further minimising heat loss to the surrounding
environment. Like penguins, the flippers of seals also have anatomical heat
exchangers in which each major artery is surrounded by a network of veins so
that heat from the former is absorbed by the latter and not lost to the
environment. During warm weather or strenuous exercise the heat exchanger
can be partially bypassed by directing more blood into surface vessels in the skin
thus enhancing the loss of heat to the environment.
Seals such as the Weddell, which has been closely studied, are consummate
divers (Kooyman, 1981). They can dive to depths in excess of 600 m and can
remain immersed for over 60 min. Several adaptations contribute to this
extraordinary capability. These include the ability to store large amounts of
oxygen in their body tissues as a result of their size, and of the relative
abundance of the oxygen-binding pigments haemoglobin (in the blood) and
myoglobin (in muscle). Weddell seals can also regulate their heart rate and their
blood flow to different organs and tissues to conserve oxygen usage during
diving. Being such excellent divers raises the question of how they avoid getting
"the bends". The bends occur when nitrogen from the air, dissolved in the tissues
under pressure, bubbles out when the pressure is reduced. It seems that seals
avoid the bends by minimising the absorption of nitrogen by partially collapsing
their lungs before diving, thus purging inhaled air and limiting its contact with the
blood. During deep feeding dives the lungs of Weddell seals collapse completely
as their thorax is crushed by the extreme pressure, but the ribs, being
cartilaginous and flexible ,are able to deform and resist breakage. Weddell seals
hunt their prey under the sea ice in limited light conditions. They have adapted to
this niche by evolving large eyes with the anatomical features typical of nocturnal
animals. An excellent summary of physiological adaptations in the Antarctic seals
is provided by Macdonald and Montgomery (1990).
Whales
A number of different whale species are found in Antarctic waters. These include
the sperm whale (Physeter macrocephalus) and the orca or killer whale (Orcinus
orca) among the toothed whales, and the sei (Balaenoptera borealis), fin
(Balaenoptera physalus), minke (Balaenoptera acutorostrata), humpback
(Megaptera novaeangliae), right (Balaena glacialis) and blue (Balaenoptera
musculus) among the baleen whales. The latter group of whales is named after
the horny plates (baleen) lined with bristles which sieve the krill and smaller
planktonic organisms on which they feed. The blue whale is the largest of all
whale species reaching 30 m in length and weighing up to 180 tonnes. The killer
whale, however, is indisputably the top predator of the Southern Ocean
ecosystem, feeding on squid, fish, seals, penguins and other whales. They can
reach 9 m in length and weigh up to 8 tonnes.
Life on the land
The Antarctic continent is host to a small number of fungal, plant and animal
species, including lichens, mosses, mites, springtails and midges. Lichens
proliferate because there is little competition from mosses, and because of their
ability to withstand drought and cold.

The Collembolan (
Gomphiocephalus hodgsoni )
This small springtail ( barely 1mm
long ) is just visible against the pale
rock background.
Gomphiocephalus hodgsoni
This scanning electron microscope
image reveals finer detail not normally
visible by eye.

The Terrestrial Food Web
On land, the primary producers which form organic material from carbon dioxide
by photosynthesis include cyanobacteria (formerly known as the blue-green
algae), diatoms, algae, lichens, mosses and liverworts. Living among them are
the decomposers, microorganisms such as filamentous fungi which feed on the
organic material made by the primary producers. Single celled protists,
microscopic animals such as rotifers and tardigrades, and larger mites and
collembolans graze on the primary producers and the decomposers.
Cyanobacteria, various microorganisms, as well as certain algae and lichens can
actually live and grow inside rocks, such as the sandstone of the Dry Valleys,
where they form what are known as "endoliths".
The ponds and lakes which are found in Antarctica usually have a benthic
community dominated by a carpet of cyanobacteria, and a planktonic community
dominated by unicellular algae.
Antarctic Fish Evolution and
Adaptation to Environment
Web Page Produced for:
New Zealand Secondary Schools
Extracts from UE, Bursaries and Scholarship Biology Prescription
Aims
1. Investigate and develop an understanding of diversity, structure, function, and
interrelationships of living organisms, and their interactions with the abiotic environment.
Objectives
Investigate and identify aspects of animal behaviour and plant responses in relation to
biotic and abiotic environmental factors.
Investigate an aspect of the ecological niche of an animal and a plant.
Investigate and describe gene expression
Investigate and explain speciation and identify patterns of evolution, with emphasis on
New Zealand examples.

Content Areas
Evolution
Adaptive radiation within one group of organisms should be studied.
As the pieces of Gondwana continued to move away from each other, the Antarctic
continent shifted to its now polar position, Drakes passage opened up and the
circulation pattern of the Southern Ocean developed. The study of the origins of the
circulation pattern of the Southern Ocean has been the subject of a recent deep sea
drilling program off the Chatham Rise. This led to Antarctica as the place we know
today, a frozen continent surrounded by ice-covered seas. What happened to the
temperate fish species that originally inhabited Antarctic coastal waters? We now know
that virtually all of these species were wiped out! One species survived probably
because of the evolution of antifreeze. Antifreeze evolved from a precursor of one of the
stomach enzymes (trypsinogen. This ancestral species founded a sub-order of fishes
known as the Notothenioidei. Because of the known time scale of the radiation of this
group, and the very cold conditions of the Antarctic seas, these fishes form a model
system for the study of issues of evolution, and adaptation to environment.
Family Tree of Antarctic Fishes


In McMurdo
Sound, the most
common
notothenioids are
the group
Nototheniidae,
these are the so-
called Antarctic
cod. The top 3
pictures shown
below are fish from
this group. They
include (left to
right) Trematomus hansoni, Pagothenia borchgrevinki, and the giant Antarctic cod
Dissostichus mawsoni. On the bottom row are members of other families, the
dragonfish Gymnodraco acuticeps, the plunderfish with the barbel on its chin, and 1 of
the icefish (Channichthyidae).


NOTOTHENIIDAE


Trematomus hansoni Pagothenia borchgrevinki Dissostichus mawsoni

Gymnodraco acuticeps Histiodraco velifer Chionodraco hamatus
White-blooded Icefishes.
The Channichthyidae have no haemoglobin in their blood, so the blood is colour-less.
Haemoglobin is an oxygen carrying pigment. Most adult vertebrates cannot survive
without its assistance. It is only the very cold waters of Antarctica which make this
possible. At some point, right at the start of the evolution of this group the ability to
make haemoglobin was lost. This loss was probably detrimental, but not lethal. We
can tell this because subsequent evolution has increased the blood volume, and the
size of the heart to make good the loss. This is one of the very few instances where we
can document the progress of adaptive evolution going backwards (a little), and then
recovering.
Antifreeze
Antifreeze is one of the most interesting adaptations to environment. Because the
blood of fishes is less salty than that of seawater it would freeze at the normal freezing
point of seawater (why is that?). This means that to stay unfrozen the fish need
antifreeze in the blood just like antifreeze is added to your car radiator to prevent
freezing. In fish the antifreeze is a glycoprotein.

Sensory Adaptations to the Antarctic Winter
The water temperature in Antarctica is right on the point of freezing all year round. It
gets no colder in the water over winter, but it is dark. In order to survive the Antarctic
winter the fish need to be able to feed in the dark. they have a specialised sensory
system called the mechanosensory lateral line, which enables them to feel the motion of
other animals in the water around them. All fish have this system, but it is particularly
well developed in deep sea fishes and in the Antarctic fish. In the picture of
Trematomus hansoni, you can see the lateral line running down the side of the body. It
consist of patches of hair cells covered by a dome. Water pushing on the dome deflects
the hair cells and this message is sent by nerves to the brain. The system exists as
superficial patches on the surface of the skin (left diagram) or patches sunk within
canals (right diagram).


The last article on this page is an abstract of a paper in the 1998 SCAR Symposium
(Antarctic Ecosystems: Models for Wider Ecological Understanding Eds W. Davison, C.
Howard-Williams and P. Broady (2000), Caxton Press. Christchurch, 2000, ISBN 0-473-
06877-X).
ADAPTIVE EVOLUTION OF ANTARCTIC FISHES
John Montgomery, School of Biological Sciences, University of Auckland, Private
Bag 92019, Auckland, New Zealand
The morphological and physiological attributes of animals change through time.
Adaptive evolution is the component of this process where selection pressures have
shaped change in particular directions. However, objectively recognizing "adaptive
evolution" as opposed to other mechanisms of change is a fraught business.
Fortunately, the Antarctic fish fauna provides an excellent system in which these issues
can be explored. Notothenioid fishes form a monophyletic radiation that dominates the
Antarctic shelf and upper slope ichthyofauna. This phylogenetic grouping, coupled with
reasonable historical re-constructions of Antarctic climate and the generally extreme
nature of the environment all contribute to the possibility of identifying important
selection forces and resulting adaptive responses. This review discusses the general
nature of the adaptations, and the confidence with which we can label particular
attributes "adaptive". It will also attempt to put aspects of the adaptive evolution of
Antarctic fishes into a comparative context. Discussions of adaptive evolution in
Antarctic fishes have centered around three areas: adaptation to low temperature; re-
invasion of pelagic habitat; and adaptation to low light. This review will pose the
questions: How do low temperature adaptations of Antarctic fish compare with low
temperature adaptations found elsewhere in Nature? What are the suite of characters
that describe pelagic fish in other oceans, and to what extent are these matched by
notothenioid pelagic species? How do the sensory systems of Antarctic fish compare
with those of deep-sea fishes? This comparative perspective provides a backdrop of
"what is possible elsewhere" against which to view the nature and the extent of the
adaptations of Antarctic fishes.
Human Involvement and Impact
Tourism
The first tourists visited Antarctica in 1958. Around 10,000 tourists now visit the
continent each Austral summer (December-February), most by ship. At issue is how to
protect the Antarctic landscape and its biota while allowing the tourists to visit and
experience Antarctica for themselves. Strict visitor guidelines are in place to minimise
potential damage, but there still remains a real threat of a significant environmental
disaster perhaps resulting from a cruise vessel going aground.
The Erebus tragedy
The first tourist flight to Antarctica was from Chile in 1956. Overflights were
introduced by Qantas and Air New Zealand in 1977 but were suspended after the
Erebus tragedy. They have since been re-established by Qantas and are still
popular (3,146 passengers in 1997-8). The Erebus tragedy involved an Air New
Zealand DC-10 on a sight-seeing flight over the Ross Sea region which crashed
into the slopes of Mt. Erebus on 28 November 1979. All 237 passengers and 20
crew were killed in New Zealands greatest air disaster.

Mount Erebus with a south polar skua
in the foreground

Political
Despite the fact that seven countries have staked territorial claims, no national
government has internationally-recognised jurisdiction over any part of Antarctica.
The Antarctic Treaty (1961)
The Antarctic Treaty was conceived in the aftermath of the very successful International
Geophysical Year (1957) which involved scientists from 67 countries. The Treaty
governs human activities in Antarctica (south of latitude 60 S) retaining it for peaceful
purposes and preserving the environment. It was originally signed in 1959 by 12
participating countries and became effective in 1961. The Antarctic Treaty was a
significant step forward in the recognition of international responsibility for the
environment, and yet it was conceived at the height of the Cold War. Since 1961 the
Treaty has been supplemented by further agreements on conservation and
environmental protection. An excellent summary of the Antarctic Treaty is provided in
May (1988).
Protocol on Environmental Protection.
The Protocol on Environmental protection was signed in 1991 and came into force in
1998. It provides for environmental impact assessment (which is required for all
activities), the conservation of flora and fauna (harmful interference of native biota is
prohibited accept by permit), the prevention of marine pollution (discharges of oils,
chemicals and garbage are prohibited), and the protection of sites of special scientific
interest (permits are required for entrance into protected areas). The Treaty also
specifies obligations with respect to waste disposal and management (abandoned
sites/dumps must be cleaned; no PCBs, pesticides), and prohibits commercial mineral
resource exploitation. Studies to date suggest that there are no large, economically
viable deposits of minerals in Antarctica, but significant mineral resources may lie
beneath the ice.
Human Exploitation of Antarctica
Antarctica has been exploited by humans since Cook first reported on the abundance of
seals and whales in the Southern Ocean.
Sealing
Sealing began in Antarctica in the early 19
th
century and only stopped when seal
numbers declined to such a low level that the operations were no longer financially
viable. The Convention for the Conservation of Antarctic Seals came into effect in 1978
providing for the total protection of fur, elephant and Ross seals south of 60 S and
setting limits to annual catches for crabeater, leopard and Weddell seals.
Whaling
In 1895 H.J. Bull organised a whaling expedition south using the Antarctica. From a
whaling perspective the voyage was far from successful, but its members did become
the first to set foot on the continent. In 1904, the Norwegian C. A. Larsen established a
whaling station at Grytviken in South Georgia which took 195 whales in its first season.
The seasonal whale kill soon reached huge proportions and the station finally closed in
1965 when the whale stocks were all but exhausted. In 1923, Larsen took a factory ship
(the Sir James Clark Ross) south and anchored it on the eastern Ross Ice Shelf where
221 whales were caught, slaughtered and processed. Very soon factory ships were
developed which had no need to anchor and the wholesale exploitation of whales was
underway. Commercial whaling is now tightly regulated and the collection of certain
species is prohibited.
Fishing
Commercial fishing in Antarctica began in the 1960s, mostly for fish but also for krill.
The dominant species in the early Antarctic fishery was the channichthyid
Champsocephalus gunnari which represented about 41.9% of the 2.7 x 10
6
tonnes of
fish caught in the first 19 years of licensed commercial operation. Fishing to this extent
has had considerable impact on fish numbers. Recent attention has focused on the
toothfish, which are now at risk of over exploitation. There are two known species of
toothfish, both of which are mid to deep water species. The Antarctic toothfish
(Dissostichus mawsoni) is found south of the Antarctic convergence while the
Patagonian toothfish (Dissostichus eleginoides) is found in sub-Antarctic waters north of
the convergence where it is fished to depths of 3500m. It reaches sexual maturity
between 10 and 12 years and lives for about 50 years, attaining a length of around 2.2
m. The Patagonian toothfish, a prized delicacy in Japan, has already been over
exploited by the fishing industry. In 1998 it was estimated that the total catch of this
species was some 10-fold over the legal catch
limit.


The Antarctic toothfish ( Dissostichus
mawsoni )


Question
Discuss problems associated with the overexploitation of natural resources.
Human Impact on Antarctica
It is important to appreciate that human activities can impact on Antarctica as the result
of both local and global actions. Fishing, for example, is a local activity whereas
damage to the ozone layer is due to the release of synthetic ozone-depleting
substances in other parts of the world. Waste from Antarctic research activities has
been a major local source of pollution in the past. The Environmental Protocol provides
for the discharge into the sea of sewage and food waste, but all other waste has to be
removed from Antarctica, usually by ship. Raw sewage contains potentially dangerous
bacteria and viruses, detergents, solvents and metals, and nitrates and phosphates
which have implications on the health of the ecosystem into which they are discharged.
Significant sewage discharges can also carpet the sea floor smothering substrate-
dwelling organisms as is seen in the vicinity of the McMurdo Station sewerage outlet.


Scott's hut at Cape Evans with Mount
Erebus in the background.



Nuclear Power
The Treaty bans nuclear explosions and nuclear waste disposal in Antarctica but does
not forbid the use of radioisotopes in science or in the generation of nuclear power.
McMurdo Base was the sight of a portable 1.8 MW nuclear power station ("Nukey Poo")
erected halfway up Observation Hill by the US Navy in 1962. The high hopes held out
for Nukey Poo never eventuated and it struggled through 10 years of operation beset
with shutdowns and radiation leaks. The reactor was eventually shipped back to the
United Sates along with 101 large drums of radioactive earth. Later, another 11,000
cubic metres of contaminated rock were removed. It took six years before the site was
decontaminated enough for unrestricted use (May, 1988). A second nuclear power plant
was planned for McMurdo Station, and others for Byrd Station, and the South Pole but
they never eventuated.
Question
What are the risks and benefits associated with nuclear power generation in
Antarctica?
Petroleum Pollution
The largest marine oil spill in Antarctica occurred on 29 January 1989 when the
Argentine supply ship the Bahia Paraiso ran aground 2 km out from the American base
of Palmer Station on the Antarctic Peninsula. A 30 m long gash was ripped open in its
hull and a 100 km
2
oil slick spread out from the vessel over the next few days. The ship
freed itself on 31 January allowing it to drift on the currents to the nearby De Laca island
where it upturned and sank to rest on the substrate with part of its keel exposed. Clean
up and containment was initially limited because of the lack of facilities at Palmer
Station, but by mid-March (after the arrival of specialist personnel from USA) about
65,000 litres of the total 1,100,000 litres on board had been collected. The impact of the
escape of fuel oils was greatest on the adjoining coastline where limpets, Adelie
penguins and blue-eyed shags were adversely affected. The effects were relatively
minor, however, and within 4-6 weeks of the spill numbers of limpets had returned to
normal (probably due to recolonisation of vacated space). A total of about 300 dead
birds were recovered, although this is probably an underestimate of the actual number
killed because of the poor weather conditions which hampered collection. The chicks of
blue-eyed shags were also adversely affected, with a number dying directly from oil
toxicity or subsequently as a result of abandonment. Most other species were
marginally affected if at all.
Question
What unique features of the Antarctic environment provide problems in dealing with oil
spills, whether minor (fractions of a litre) or catastrophic (millions of litres) in nature?
Impact of Research Stations
Although most of Antarctica is relatively pristine, high levels of pollutants have been
recorded in localised areas such as in the vicinity of scientific bases like McMurdo
Station (reviewed in Lenihan et al., 1992). McMurdo Station is the largest human
settlement in Antarctica, with a summer population of approximately 1000 people. Since
construction of the base about 40 years ago, poor waste discharge practices have
resulted in an intense contamination gradient of polycyclic aromatic hydrocarbons
(PAHs) and polychlorinated biphenyls (PCBs) in sediments of the adjacent Winter
Quarters Bay. Sediment levels of hydrocarbons as high as 4.5 g kg
-1
have been
detected in the upper reaches of the bay. This is higher than levels seen in the most
polluted harbours of other parts of the world. A similar pattern has been reported for
PCBs, with levels as high as 1.4 mg kg
-1
in the upper reaches of Winter Quarters Bay,
and a marked change in marine benthic communities along the contamination gradient
has been noted. A large amount of anthropogenic debris, such as used machinery, 44
gallon drums and scrap metal is also to be found littering the sea floor in Winter
Quarters Bay.


The American research base McMurdo Station located on Ross Island.



An extensive clean up program was begun at McMurdo Station in 1988. Major rubbish
dumps were removed and returned to the USA, dumping along the shoreline was
prohibited, and sewage and grey water were required to be discharged at a submerged
outfall. The results of a recent study by Miller et al. (1999), however, show that a
significant level of contamination still exists in Winter Quarters Bay, and that it has the
potential to impact on resident organisms such as the local fish species Trematomus
bernacchii. Since hydrocarbon breakdown is slow in Antarctica due to the cold
temperatures, the effects of earlier contamination episodes are likely to remain for a
considerable period of time. Is this to be the legacy we leave future Antarctic visitors?


The New Zealand research station Scott
Base.