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13, 1986). The intensity of summer populations. This structure is perhaps from food, much of which is recycled
production is a major reason why surprising given that krill are moved in the water column as it sinks to the
this area, comprising just 10% of huge distances by the powerful seabed.
the global ocean, is responsible Antarctic Circumpolar Current. Ecologists have long been
for~20% of CO2 draw-down. There Perhaps the most striking feature of intrigued as to whether the extreme
are many species of algae involved the water column biota of the Southern physical conditions of the polar
and the composition of blooms vary Ocean is the paucity and low diversity oceans, the unusual nature of the
with water mass and with proximity of fish. A few groups, most notably Antarctic continental shelf and/or its
to oceanic fronts or the seasonal the notothenoids, have radiated in isolation by the Antarctic Circumpolar
sea-ice margin. In consequence, Antarctica but many taxa and guilds Current have resulted in a fauna
the composition of phytoplankton familiar from lower latitudes are that differs from that elsewhere.
preserved in sediment cores has completely absent. The notothenioids The Southern Ocean is the only
proved to be a valuable tool for dominate biomass and diversity marine region in which sharks and
investigating historical variation in on the continental shelves around turtles are absent, and likewise
oceanography, such as where and Antarctica. They are a predominantly there are benthic groups which are
how far fronts move northwards benthic group, having evolved form an absent (for example, brachyuran
during ice-ages. ancestor that had lost the function of crabs) or very rare (such as acorn
The phytoplankton biomass is the swimbladder. They tend to be slow barnacles, rays and reptant decapod
dominated by large diatom species, moving, and relatively few species have crustaceans). However, the benthos
though some areas can be dominated evolved the ability to live in the water is just as remarkable for those
by Phaeocystis and many consumers column. Their physiology has attracted groups that are well represented;
may target the smaller ciliates and a great deal of attention, particularly pycnogona (sea spiders), peracarids
flagellates (nano-phytoplankton) or in relation to freezing resistance and (amphipods and isopods) and
bacteria (picophytoplankton). The metabolism. Because teleost (bony) bryozoans are all unusually rich
interaction and lag between physical fish have blood that is more dilute than and abundant. In common with
variability and changes in primary seawater, they run the risk of freezing most isolated regions,most
production has been a key area of in very cold polar waters. Freezing of these species appear to be
research, particularly with respect is prevented by the presence of a endemic. Debate continues
to its trophic cascade effect on glycoprotein antifreeze; this prevents as to which factors havebeen
zooplankton and their high predators. small ice crystals from growing to a most important inproducing
The fate of most primary production dangerous size, though how the fish this diverse and unusualfauna.
in the Southern Ocean is to be broken clears this ice from its system is still an Recent suggestions include lack of
down by bacteria and protists in the area of active research. One group of predators able to crush skeletons,
microbial web, and new molecular notothenioids, the ice-fish, no longer the unusuallysediment-free water
approaches are revealing huge express the genes for haemoglobin column, to which can be added the
and previously unrealised bacterial (and often myoglobin) and so their long-running theme of history.
biodiversity. A second important blood is clear and translucent. At the More than 9700 species are
route for algal productivity in the low temperatures of polar waters these described from the Southern Ocean,
Southern Ocean is consumption by fish can carry sufficient oxygen for most of which are benthos, but we
the most numerous large animals in metabolism in simple solution. Most know much less about them than
the world: Antarctic krill (Euphausia Antarctic fish grow slowly and do the pelagos (Figure 2). It had long
superba) and calanoid copepods. not reach maturation for many years, been thought that species richness
Krill can form swarms more than a making them highly vulnerable to in general declined pole-wards. This
km long; they grow through a series harvesting. Although compared with is clearly the case on land and in
of moult stages to reach more than elsewhere Southern Ocean fisheries freshwater, but although it is true for
five centimetres in length and annual have been fairly small scale, targeted macro-algae and a few animal groups
production is estimated to be 340540 populations have still not recovered (notably gastropod molluscs) in the
million tonnes. Zooplankton, and decades later. sea, other marine taxa are as rich
especially krill, are the key prey for or richer than at low latitudes (for
most of the worlds penguins and A deep but diverse sea-bed example, holothurian sea cucumbers
baleen whales, as well as many fish, Water column-seabed interactions or ascidian sea squirts). The diverse
squid, seabirds and seals. Krill is also (benthopelagic coupling) may be Antarctic seabed thus contrasts with
the target of a substantial fishery, at its weakest in high latitudes. patterns on land, freshwater and in
and so understanding its population This is because the vast icesheet the water column, and indeed it is
dynamics has long been seen as a on the continent of Antarctica has comparable in diversity across all
priority for science and conservation depressed the shelf so it is deeper taxonomic levels with all but coral
in the region. Recent genetic than elsewhere, and together with reef ecosystems. For example, more
studies have shown that clock the scouring of the shelf by previous than 1200 species are known from
genes genes for components of extensions of the ice sheet at the South Orkney Islands and nearly
the molecular machinery controlling glacial maxima this has resulted in 2000 around South Georgia. The
circadian rhythms control moulting a continental shelf much deeper number of known benthic species
and diel migration, and other work than the photic zone of primary has near doubled in the last decade,
has suggested that there is weak but production. In consequence, benthic partly as a result of particularly
significant geographic structure to primary consumers are further away intense multinational efforts during
Current Biology Vol 21 No 12
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50
50
areas as big as the Amundsen Sea,
S
S
which is 40 longitude wide, were
60
60
S
S
only first sampled in 2008) but also 60E 60E
70
S
The Southern Ocean has been
80
80
claimed to be the most isolated of
S
any large marine area because it 90W 90E 90W 90E
juveniles or young; communities are A considerable literature has already Rising surface temperatures have
in various stages of recovery from been built up on ectotherm responses also led to much speculation about
scouring by icebergs. Most can to acutely raised acidification through invasions of the Antarctic continental
recruit from deeper water as Antarctic laboratory experimental manipulations shelf by non-indigenous species or
benthos occur across unusually wide on various life history stages of a anomuran (stone) crabs from deeper
depth ranges (eurybathy), a feature range of Antarctic taxa. These suggest water with consequent changes
which may be related to previous differing degrees of tolerance (that is, to assemblage structure or even
extensions of the ice sheet driving the ability to maintain shell thickness), extinctions of native fauna. Whilst
many species off the shelf and in to but the extent to which any individual such speculation has attracted
deeper water. species will cope through tolerance, considerable attention, hard evidence
Many benthic species recorded as against adaptation or migration, is of actual biological change with
from the shallows have ranges unknown and hard to test over realistic a clear causal link to changes in
beyond the shelf break at 800 m and time scales. One of the key problems climate is essentially confined
down the continental slope. Very little of assessing in situ biological response to changes in the distribution of
is known about the biology of the to acidification is the paucity of historic phytoplankton and sea-bird (mainly
continental slope and deep sea, even measurements of pH, something which penguin) distribution. Thus, even
in the best-studied regions such as is now being addressed by examination in an area experiencing one of the
the Weddell Sea, although a recent of museum specimens with skeletons fastest rates of regional climate
series of scientific cruises (Antarctic of differing carbonate structure. change, and despite considerable
Deep sea or ANDEEP) have shown Much experimental work has international marine biological effort,
that some groups, notably isopods, also been directed at the response the biological response to physical
are unusually rich here. Whether most of nearshore benthos to acute change is not yet clear. We suggest
abyssal species are rare or intensely temperature rises, showing that many that impacts are most likely to be
patchy has become a source of common species in the shallows are detected near research stations
debate, as has been the scalability unable to perform critical activities in the shallower waters of the
of richness estimates (that is, how following increases of just 23C. As Bellingshausen and Scotia shelves.
valid is it to scale up from the results with studies of the acute response These will probably involve one or
of a few samples to the deep-sea to acidification, it is difficult to more of the first detection of range
environment as a whole?). Except assess relevance of short-term shifts in the more sensitive species,
for meiofauna, only in the last few studies to understanding the increased growth by suspension
years has sampling protocol in deep long-term responses to environmental feeders, differential survival across
waters started to become designed change. Arguably many of the most age spectra of well monitored
more critically to increase statistical likely marine biological responses species, arrivals of non-indigenous
power. The last decade has marked will be through altered community species, and increased ice-scour
a step change in the development of interactions, but in situ study of this disturbance in the shallows (as
collecting, describing and databasing is greatly complicated by iceberg reduced periods of winter fast ice
deep biodiversity, but the drivers activity, slow growth and community allow ice bergs to move around
of Antarctic abyssal biodiversity development, and taxonomic more).
remain obscure. Elucidating these resolution.
may be crucial to reconstructing One consequence of the Marine biology as a tool to interpret
and understanding Antarcticas widespread retreat of glaciers and wider science questions
past and will require closer linkage ice-shelves is that phytoplankton Historically marine biology
across disciplines, as has happened blooms now occur in areas previously in Antarctica has benefitted
during the latest exploration visits covered by shelf ice, and also greatly from synergy with other
to cold seep and hydrothermal vent further south where the duration disciplines, particularly geology and
communities in Antarctica. of winter sea ice has shortened. oceanography, but also geography,
Oceanographic changes to the west glaciology and meteorology, all of
Marine biological response to of the Antarctic Peninsula have also which provide valuable context for
physical change affected higher levels in the food- the interpretation of biological data.
The ocean to the west of the Antarctic web. A decrease in krill biomass A multidisciplinary approach to the
Peninsula is warming, and projections and a concurrent increase in salps testing of biological hypotheses is
from global models suggest that over the last few decades has been now widespread, but in recent years
this will be amongst the first places reported, though links to climate biological data have proved critical
to become under-saturated first in change remain speculative. More to other disciplines. This is because
aragonite and then in calcite; in other clearly linked to warming have been it can provide data where there are
words, this is a likely hotspot of ocean marked changes in the distribution otherwise little or none, or it can
acidification. Measurements to date of penguin breeding colonies, with provide improved spatial or temporal
already show that near surface waters populations of the ice-associated resolution.
of the Bellingshausen and Scotia seas Adlie penguin having decreased Perhaps the best recent example
have warmed as fast as anywhere in towards the north of their range and has been the use of biological
the last 50 years, and that the highest increased at more southerly locations distributions to refine models of ice
rate of glacier retreat, ice shelf and and an associated shift southwards sheet behaviour under past climate
sea ice losses are along the Antarctic of the related Gentoo and Chinstrap variation. Ice cores provide a high
Peninsula. penguins. resolution record of past atmospheric
Current Biology Vol 21 No 12
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Concluding remarks
Marine biology was one of the
earliest scientific disciplines to be
undertaken in Antarctica. Whilst
this led to a rapid improvement in
our understanding of the Southern
Ocean and its biota, it also led to an
explosion of resource harvesting.
Early exploitationfocussedon
whales and seals, but since the
1960s there have also been finfish
and krill fisheries. The Antarctic
Figure 3. Schematics of Antarctica, the Southern ocean sea-floor and ice extent now and as also has a long and distinguished
hypothesised in one of the last interglacial periods. history of fundamentalresearch on
The huge ice sheets weigh down Antarctica and its surrounding continental shelf (yellow), many of these exploited species,
making it the deepest (as well as the widest) globally. Geological and biological data suggest research which has not only given
species in the Weddell and Ross Sea, which are separated by the West Antarctic Ice sheet,
were connected perhaps as recently as 120,000 years ago. Studying signals of connectivity
us fundamental insights into their
in species of the region could help reconstruct ice sheet behaviour during historic periods of population dynamics and the
warming and thus more accurately model future sea level change. We thank Peter Fretwell for relationship of these dynamics to
help making this figure. large-scale oceanographic and
atmospheric phenomena such as
temperature and composition but on nearly all shelf. However, recent Southern Annular Mode and the El
these data are spatially limited. careful biogeographic work has Nio Southern Oscillation (ENSO),
Glaciologists and geographers have suggested that the Weddell and but also provided the first signals
mapped ice sheet topography and Ross seas, on either side of the West ofpopulation decline (for example,
thickness and can determine ice Antarctic ice sheet, were connected inmany seabirds which were
thinning patterns and the topography recently. Such a marine connection caughtas by-catch in fisheries,
of underlying bedrock to reveal likely suggests a partial collapse of the both within the Southern Ocean and
paths of weakness and instability. thirdlargest ice mass on Earth elsewhere).
However, recent work on the genetics (Figure3). This important synergy For decades, biological effort was
and distributions of terrestrial biota between biological and earth sciences largely confined to major expeditions
has shown clearly that some land is being further used to help elucidate or the immediate environs of research
has remained ice-free; this then the pathand extent of the seaways stations serviced by ship. However,
constrains model reconstructions of and their timing, which should give the development of permanent
past ice-sheet size and distribution. strong clues to West Antarctic ice stations with air-bridges, and
On the continental shelf, studies of sheet behaviour in past interglacial the greater use of new sampling
ice scouring lines, moraines and periods and how global sea level is techniques such as SCUBA and
sediments suggest that at the last likely to change under current and remotely operated vehicles have
glacial maximum, ice sheets grounded projecting warming. greatly increased both the spreads
Magazine
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