30° 0° 30°

CONVER
IC— GE
R CT NC
E
TA
N
A

Prince Edward
Bouvetaya Marion
Island
Island
South South Sandwich
Georgia Islands
60° lies 60°
Crozet

Falkland South Orkney Sanae

Neumayer
SOUTH Islands Islands
AMERICA
Signy Syowa
South QUEEN MAUD
Shetland 75°S LAND lies
Islands Larsen Halley Kerguelen
ENDERBY McDonald Is
Ice Shelf LAND Mawson
AN Ronne Heard
T Island
Ice Prydz Bay
AR

Shelf GREATER
TIC

ANTARCTICA Davis
Siple
PE
NI

90° LESSER Amundsen Mirny 90°

NS

Scott
UL

Peter I Island ANTARCTICA

A

Oasis
WILKES
MARIE BYRD LAND
Ross Casey
LAND
Ice Shelf
VICTORIA
ROSS LAND
SEA
McMurdo Scott
75°W 70° 65° 60° 55°W
Bellingshausen Hallett
Frei Dumont
Balleny D'Urville
Arctowski
Deception I Islands
Scott Island

Palmer
65°S Faraday
60°S 120°
Rothera Almirante
Brown Macquarie
Island
Campbell
Island
Auckland Island

70°S Fossil 45°S

Bluff
NEW AUSTRALIA
80°W ZEALAND
50°W 180° 150°

Antarctica is both a continent and an ocean. The continent is inextricably linked to the vast extent of the Southern Ocean
surrounding it, an ocean that supports a teaming array of life, ranging from microscopic plants that form part of the
phytoplankton responsible for primary production to the mighty whales, our largest marine mammals. Whilst our knowledge
of the biology of these waters has been accumulating since the first voyages of discovery in the 18th century, there is yet
to be a comprehensive, single-author synthesis of the current state of that knowledge. This book sets out to correct that
deficiency.

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Preface
Our knowledge of the biology of the seas surrounding the
Antarctic continent began with descriptions and specimens
brought back by early naturalists on voyages of discovery in
the Southern Ocean. Prominent among these were the early
French navigators such as Bouvet de Lozier, who discovered
Bouvet Island in 1739, and Yves Joseph de Kerguelen-
Tremarez, who discovered the island that bears his name
some 30 years later. These were followed by the voyages of
Captain Cook, who, in 1772, reached 718 10 0 S in the
Bellingshausen Sea. In 1819, the Russian explorer Admiral
Bellingshausen was first to tow a net at the stern of his ship
and he found that it collected more organisms during the
night than during the day. He was probably the first to collect
the Antarctica krill, Euphausia superba.
The first serious scientific marine research was carried
out by James Eights who sailed in 1929 with the American
sealing Captains Palmer and Pendleton. Eights described the
natural history of the South Shetland Islands. His best known
discovery was of a ten-legged pycnogonid. Soon after, there
followed a series of national expeditions led by Dumont
D’Urville of France (1837–1840), Charles Wilkie of the
United States (1838–1842), and Sir James Clark Ross of
Britain (1939–1943). Among the naturalists who accom-
panied these expeditions, the most notable were J.D. Dana,
on the Wilkie expedition, and J.D. Hooker, on the Ross
expedition. These expeditions all made extensive collections
of marine organisms.
There was a renewal of Antarctic exploration at the end
of the nineteenth century. Major expeditions included the
British (1899–1900), the Belgian (1898), the Swedish
National Expedition (1902–1904), the Scottish National
Antarctic Expedition (1902–1904), the German expedition
(1901–1903) in the Gauss, and the French Expedition in the
Pourquoi Pas (1908–1910). All of these expeditions resulted
in taxonomic studies that laid the foundation for later studies
of the marine flora and fauna.
Ecological studies in the Southern Ocean began with the
work of naturalists attached to land-based, over-wintering
parties such as those of Sir Ernest Shackleton’s 1907–1909
expedition and Captain Scott’s two polar expeditions.
Antarctic marine studies received a tremendous impetus
from the investigations that began in 1925 with the study of
whale carcasses at the whaling stations at Grytviken in South
Georgia. The work of the scientists expanded to include, not
only studies on whale demography and ecology, but also of
the physical, chemical and biological oceanography of the
Southern Ocean in order to gain an understanding of the
factors influencing the distribution, reproduction, and growth
of whale stocks. These investigations contributed to the first
long-term studies of the Antarctic pelagic ecosystem, and
they were the first that extended right around the Antarctic
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continent. The voluminous reports of these Discovery
expeditions laid the indispensable foundation for subsequent
studies.
Southern Ocean marine research entered a new phase
following the establishment, in 1957, of the Special (later the
Scientific) Committee on Antarctic Research (SCAR). This
initiated the modern era of Southern Ocean research in which
many countries played roles, notably the Soviet Antarctic
Expeditions in the Vitez and Ob and the United States Eltanin
cruises. Other countries such as Great Britain, France, South
Africa, Chile, Argentina, Australia and Japan were also very
active during this period. In 1972, a coordinated international
research program was initiated with the establishment of the
BIOMASS (Biological Investigations of Antarctic Systems
and Stocks) Program (El-Sayed 1996).
My own involvement in Antarctic research began with
a visit to McMurdo Sound in 1960 where one of my staff
in the Department of Zoology at the University of
Canterbury, Dr. Bernard Stonehouse, was initiating a
research program on Adelie penguins and McCormick
skuas. This led to the establishment of the University of
Canterbury Antarctic Research Unit (Knox 1986, 1988)
that carried out continuous summer research in the
McMurdo Sound region over the period 1960–1983, the
last twelve years of which I was director of the unit.
During the summers of 1969 and 1970, I initiated an
inshore marine research program that continued until the
1982–1983 season. Throughout the operation of the unit,
some sixty research students participated in its activities.
Interaction with these enthusiastic young minds had a
significant influence on many of the concepts developed in
The Biology of the Southern Ocean. I also have had the
good fortune to become involved in international Antarctic
science activities, first as a member and secretary of the
SCAR Biology Working Group on Biology and as a
member of SCAR since 1969. I was a member of the
Group of Specialists on the Living Resources of the
Southern Ocean from its inception and I attended the first
two meetings of CCAMLR (Convention on the Conserva-
tion of Antarctic Marine Living Resources). These
activities brought me into contact with a wide range of
Antarctic scientists from all the SCAR countries.
The suggestion for The Biology of the Southern Ocean
was made by Dr. Bernard Stonehouse and, without his encour-
agement, it would not have been written. Discussions with
Antarctic colleagues throughout the SCAR community have
had a profound influence on its development. Amongst these I
particularly wish to thank Professor Sayed El-Sayed, the late
Sir George Deacon, Professor A.L. DeVries, Professor M.
Fukuchi, Sir Martin Holdgate, Professor T. Hoshiai. Dr. J.C.
Hureau, Dr. K.B. Kerry, Dr. Y. Naito, the late Dr. T. Nemoto,
Dr. J. Oliver, Dr. J. Warham, and Professor E.C. Young.
Access to literature was greatly facilitated by periods spent in
the libraries of the British Antarctic Survey and the National
Institute of Polar Research, Tokyo. Dr. D.W.H. Walton
reviewed all chapters of the original edition in various drafts
and I am grateful for his assistance and encouragement. I
would also like to thank Dr. Maria Murphy and the staff of
Cambridge University Press for their patience and support
during the preparation of the first edition and for seeing the
project through to completion.
The first edition of The Biology of the Southern Ocean
was published in 1993 and was well received. The print run
was sold out a number of years ago. Some 13 years have
passed since then and a considerable amount of research has
been carried out during this period. The original SCAR
countries engaged in marine research in the Southern Ocean
have been joined by other countries including China,
Ukraine, Uruguay, Korea, Ecuador, India, Peru and Spain,
resulting in a much increased research effort.
The last decade in particular had seen the initiation of
several scientific programs to study phenomena and
processes of global significance in which the Southern
Ocean plays a key role. Some of the major programs are
listed below and brief descriptions of the objectives of these
are given in El-Sayed (1991).
- Antarctic Marine Living Resources (AMLR) program
(1986-present)
- Antarctic Marine Ecosystem Research at the Ice-Edge
Zone (AMERIEZ) (1983–1988)
- Research on Antarctic Coastal Ecosystem Rates
Program (RACER) (1986–1993)
- Long-Term Ecological Research (Palme LTER)
(1980-present)
- Southern Ocean Joint-Global Ocean Ecosystem
Dynamics (SO-GLOBEC) (1991–2009)
- Coastal and Shelf Ecology of the Antarctic Sea-Ice
Zone (CS-EASIZ) (1993–2004)
- Antarctic Pack-Ice Seals (APIS) (1995-present)
- European Polarstern Study (EPOS) (1988–89)
- Research on Ocean-Atmosphere Variability and
Ecosystem Response in the Ross Sea (ROAVERRS)
(1996–1998)
The multi-disciplinary, multi-national cruises of the R/V
Polarstern have been of particular importance in recent
years, especially the ANDEP cruises (2002–2005) that
investigated the benthic fauna of the deep Weddell Sea.
Over the period 2000–2002, a series of iron fertilization
experiments (SOIRE, SOFex and EISENEX) investigated
the potential role of iron in controlling algal production in the
Southern Ocean. Recently, the New Zealand National
Institute of Water and Atmospheric Research (NIWA)
investigated the richness and diversity of the benthos in a
latitudinal gradient in the western Ross Sea in relation to
broader-scale environmental factors and local-scale variation
in productivity sources.
As the result of the research activities listed above, there
has been a considerable volume of published research on the
biology of the Southern Ocean in the 13 years since the
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publication of the first edition of The Biology of the Southern
Ocean. While some of the information in the first edition is
retained, it is essentially a new account. The task of
summarizing the large volume of research has proved a
difficult task. Most of the figures in the text have been
redrawn from the originals.
The book commences with a description of the physico-
chemical environment of the Southern Ocean and then
follows a logical sequence covering phytoplankton and
primary production, the sea ice microbial communities, and
the secondary consumers, the zooplankton. There is an
extended chapter on the biology and ecology of Antarctic
krill, in view of its central position in the Southern Ocean
food web. Additionally, krill has been the subject of intensive
research programs over the past decades, especially during
the BIOMASS Programmes and subsequently by CCAMLR.
A series of chapters consider the higher consumers, nekton
(with an emphasis on cephalopods), fish, seals, whales and
seabirds. A series of chapters then follow on selected
ecosystem components: the benthic communities, life
beneath the fast ice and ice shelves, recent advances in
understanding decomposition processes, and the role of
bacteria and protozoa. These are followed by an attempt at a
synthesis of ecosystem dynamics, with an emphasis of the
pelagic ecosystem and then three chapters dealing with
resource exploitation, the impact of such exploitation on the
marine ecosystem, and the problems involved in the
management of the living resources. Three new chapters
have been added to the second edition exploring the impact
of increased UV radiation, human impact on the marine
environment, and the impact of global warming on Southern
Ocean marine ecosystems.
As can be seen by the reference list at the end of this
volume, there is a considerable body of recent literature on
the biology of the Southern Ocean. Because of the growth of
published research, I have had to be selective in the material
that is included. Examples have been carefully chosen from
the pool of published research to illustrate the concepts
discussed. There are, doubtless, others that could have been
used and I apologize to authors whose work has not been
included.
In the preparation of this volume I have been supplied
with advanced copies of papers prepared or accepted for
publication and I am grateful to all who supplied them. I am
also appreciative of the support and encouragement of John
Sulzycki and the staff of CRC Press, in particular Pat
Roberson, during the preparation of this edition and for
seeing the project through to the completion of a high quality
product.
This book is an attempt to synthesize the available
information into a coherent account of one of the most
fascinating systems on the globe. I hope that it will prove
useful to advanced undergraduates and to professionals
engaged in Antarctic marine research, as well as to all
interested or involved in Antarctic marine conservation and
management. It will have been worthwhile if it stimulates
others to work on the fascinating aspects of Antarctic marine
research that have been discussed in this book.
Acknowledgments
I would firstly express my indebtedness to the many international Antarctic scientists, some of whom have been listed in the
Preface, who I have interacted with over the years. Discussions with them have influenced the development of many of the
concepts dicussed in this book. I acknowledge the outstanding contribution of Cambridge University Press, in the preparation
of the first edition of this book. I owe a great debt to the staff members and students who participated in the research activities of
the University of Canterbury Antarctic Research Unit.
I owe a great deal to John Sulzycki and the staff of CRC Press, in particular Pat Roberson for their patience and support
during the preparation of this second edition. Together with the staff of Alden Press Services who typeset this book they have
produced a high quality product.

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Author
George A. Knox, C.N.Z.M., M.B.E., F.R.S.N.Z., was head
of the Department of Zoology, University of Canterbury,
Christchurch, New Zealand, from 1959 to 1976. He is now
professor emeritus of zoology.
Professor Knox was born in New Zealand and received
his education at the University of Canterbury, where he was
appointed a staff member in 1948. He has been a visiting
fellow at the East-West Center, Honolulu, and a visiting
professor at the Department of Oceanography, Texas A&M
University and the Department of Environmental Engin-
eering, University of Florida, Gainesville. He has visited and
worked in laboratories in the U.S., Canada, Chile, Japan,
Australia, Western Europe, the USSR, and China.
Professor Knox’s research has been wide ranging and
includes: (1) the systematics and distribution of polychaeta
with special reference to New Zealand and Antarctica; (2)
rocky shore intertidal ecology and biogeography; (3) the
ecology and conservation of islands; (4) studies on the pelagic
and benthic ecosystems beneath the sea ice in McMurdo
Sound, Antarctica; and (5) estuarine and coastal ecology and
management. He established and directed the Estuarine
Research Unit in the Department of Zoology and the
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University of Canterbury Antarctic Research Unit. He
has participated in many field expeditions, including the
Chatham Islands 1954 Expedition (leader); the Royal Society
of London Darwin Centennial Expedition to southern Chile
(marine biologist and deputy leader); thirteen summer
expeditions to McMurdo Sound, Antarctica; the establishment
of the Snares Islands Research Program (he participated
in three field expeditions); and participation in field
expeditions to Campbell and Auckland Islands. He has
published over 100 scientific papers and 28 environmental
reports, written five books, and edited and co-authored three
other volumes.
Professor Knox has received a number of awards and
fellowships for his contributions to science, including Fellow
of the Royal Society of New Zealand (FRSNZ), 1963; Hutton
Medal, Royal Society of New Zealand, 1978; Conservation
Trophy, New Zealand Antarctic Society; Honorary Member of
Scientific Committee on Antarctic Research, 1982; member of
the Most Excellent Order of the British Empire (MBE), 1985;
the New Zealand Association of Scientists’ Sir Ernest Marsden
Medal for Service to Science, 1985; and Companion of the
New Zealand Order of Merit (CNZM), 2001.
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Contents
Chapter 1 The Southern Ocean
1.1 Introduction.................................................................................................................................................. 1
1.2 The Evolution of the Southern Ocean......................................................................................................... 3
1.3 Bathymetry................................................................................................................................................... 4
1.4 Climate......................................................................................................................................................... 4
1.5 Ice Cover...................................................................................................................................................... 6
1.6 Circulation Patterns and Water Masses ...................................................................................................... 7
1.7 Some Regional Hydrographic Features..................................................................................................... 13
1.8 Bottom Water Formation........................................................................................................................... 14
1.9 Nutrients..................................................................................................................................................... 15

Chapter 2 Phytoplankton and Primary Production

2.1 Introduction................................................................................................................................................ 18
2.2 Data Base ................................................................................................................................................... 19
2.3 Species Composition and Distribution...................................................................................................... 19
2.4 Latitudinal Zonation of the Planktonic Subsystems ................................................................................. 24
2.5 Phytoplankton Biomass ............................................................................................................................. 30
2.6 Primary Production.................................................................................................................................... 32
2.7 Seasonal and Geographic Variation of Phytoplankton Biomass and Primary Production ...................... 36
2.8 Factors Affecting Primary Production....................................................................................................... 38
2.9 Growth Rates ............................................................................................................................................. 53
2.10 Heterotrophic Nutrition ............................................................................................................................. 56
2.11 A Model of Phytoplankton Production ..................................................................................................... 56

Chapter 3 Sea-Ice Microbial Communities

3.1 Introduction................................................................................................................................................ 59
3.2 Sea Ice as a Habitat ................................................................................................................................... 60
3.3 Sea-Ice Micro- and Meiofauna.................................................................................................................. 94
3.4 Dynamics of Sea-Ice Microbial Communities.......................................................................................... 96

Chapter 4 Zooplankton
4.1 Introduction.............................................................................................................................................. 100
4.2 Species Composition and Distribution.................................................................................................... 101
4.3 Life History and Growth ......................................................................................................................... 111
4.4 Vertical Migration ................................................................................................................................... 114
4.5 Swarming ................................................................................................................................................. 115
4.6 Feeding..................................................................................................................................................... 116
4.7 Biomass and Production .......................................................................................................................... 121
4.8 Ecophysiology.......................................................................................................................................... 123
4.9 Strategies for Winter Survival................................................................................................................. 124

Chapter 5 Krill
5.1 Introduction.............................................................................................................................................. 128
5.2 Species of Euphausiids ............................................................................................................................ 129
5.3 Life History and Growth ......................................................................................................................... 129
5.4 Krill Aggregations ................................................................................................................................... 139
5.5 Distribution and Abundance.................................................................................................................... 147
5.6 General Distribution Patterns .................................................................................................................. 150
5.7 Factors Affecting Distribution................................................................................................................. 152

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 5.8 Overwintering Mechanisms..................................................................................................................... 158
5.9 Stock Seperation ...................................................................................................................................... 161
5.10 Feeding and Energy Expenditure ............................................................................................................ 162
5.11 Krill Production ....................................................................................................................................... 172

Chapter 6 Nekton
6.1 Introduction.............................................................................................................................................. 177
6.2 Species Composition and Distribution.................................................................................................... 177
6.3 Cephalopods............................................................................................................................................. 178

Chapter 7 Fish
7.1 Introduction.............................................................................................................................................. 185
7.2 Species Composition and Distribution.................................................................................................... 187
7.3 Morphological and Physiological Adaptations ....................................................................................... 191
7.4 Reproduction and Growth ....................................................................................................................... 193
7.5 Age, Growth, Mortality and Biomass Structure ..................................................................................... 196
7.6 Feeding Ecology ...................................................................................................................................... 198
7.7 Factors Controlling the Distribution, Abundance, and Trophic Ecology of Antarctic Fish.................. 207

Chapter 8 Seals
8.1 Introduction.............................................................................................................................................. 209
8.2 Species Composition and Distribution.................................................................................................... 210
8.3 Abundance ............................................................................................................................................... 215
8.4 Social Organization and Reproductive Behaviour.................................................................................. 216
8.5 Feeding Ecology ...................................................................................................................................... 220
8.6 Reproduction, Growth, and Development............................................................................................... 227
8.7 Population Dynamics............................................................................................................................... 228

Chapter 9 Whales
9.1 Introduction.............................................................................................................................................. 231
9.2 Species Composition ............................................................................................................................... 232
9.3 Distribution, General Life Histories, and Migrations ............................................................................. 235
9.4 Segregation .............................................................................................................................................. 238
9.5 Reproduction............................................................................................................................................ 239
9.6 Feeding Ecology ...................................................................................................................................... 241
9.7 Bioenergetics............................................................................................................................................ 243
9.8 Population Dynamics............................................................................................................................... 245
9.9 Role in the Ecosystem ............................................................................................................................. 247

Chapter 10 Birds
10.1 Introduction.............................................................................................................................................. 249
10.2 Species Composition and Distribution.................................................................................................... 250
10.3 Breeding Biology..................................................................................................................................... 256
10.4 Nonbreeding Biology............................................................................................................................... 259
10.5 Food and Feeding Ecology...................................................................................................................... 259
10.6 Energetics................................................................................................................................................. 270
10.7 Population Structure and Dynamics........................................................................................................ 271
10.8 Current Status and Population Trends in Antarctic Seabirds ................................................................. 272
10.9 Role in the Ecosystem ............................................................................................................................. 273

Chapter 11 Benthic Communities

11.1 Introduction.............................................................................................................................................. 275
11.2 The Antarctic Benthic Environment ....................................................................................................... 276
11.3 Littoral Communities............................................................................................................................... 277
11.4 Shallow Sublittoral Communities ........................................................................................................... 281
11.5 Epifaunal Communities ........................................................................................................................... 288

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 11.6 Infaunal Communities ........................................................................................................................... 294
11.7 Shelf and Upper Slope Communities.................................................................................................... 300
11.8 Deep Sea Communities ......................................................................................................................... 300
11.9 Microbial Communities ......................................................................................................................... 301
11.10 Meiofauna .............................................................................................................................................. 301
11.11 Biogeography and Origin of the Benthic Biota .................................................................................... 302
11.12 Factors Responsible for Shaping the Antarctic Benthos ...................................................................... 303
11.13 Biogeographical Schemes for the Antarctic Region............................................................................. 307
11.14 Diversity, Abundance, and Biomass ..................................................................................................... 307
11.15 Food and Feeding .................................................................................................................................. 309
11.16 Bentho-Pelagic Coupling....................................................................................................................... 311
11.17 Ecological Strategies ............................................................................................................................. 312

Chapter 12 The Fast Ice and Ice Shelves

12.1 Introduction............................................................................................................................................ 317
12.2 The Coastal Fast Ice Environment ........................................................................................................ 317
12.3 Primary Production................................................................................................................................ 319
12.4 Sedimentation and Resuspension .......................................................................................................... 324
12.5 Zooplankton ........................................................................................................................................... 324
12.6 The Sympagic Community.................................................................................................................... 327
12.7 The Platelet Ice Community.................................................................................................................. 330
12.8 The Cryopelagic Community ................................................................................................................ 330
12.9 The Ross and McMurdo Ice Shelves .................................................................................................... 332

Chapter 13 Ice-Edge Processes

13.1 Introduction............................................................................................................................................ 341
13.2 The Ice-Edge Habitat............................................................................................................................. 342
13.3 Ice-Edge Phytoplankton Biomass and Primary Production.................................................................. 343
13.4 Potential Causes of Phytoplankton Blooms .......................................................................................... 347
13.5 A Model of Ice Edge Bloom Dynamics ............................................................................................... 350
13.6 Bacterioplankton .................................................................................................................................... 351
13.7 Ice-Edge Microheterotrophs .................................................................................................................. 351
13.8 Ice-Edge Zooplankton and Nekton ....................................................................................................... 352
13.9 Ice-Edge Vertebrates ............................................................................................................................. 355
13.10 The Importance of the Ice Edge in the Ecology of the Southern Ocean............................................. 357

Chapter 14 Decomposition and the Roles of Bacteria and Protozoa

14.1 Introduction............................................................................................................................................ 361
14.2 Quantities and Sources of Organic Matter............................................................................................ 362
14.3 Sedimentation of POM .......................................................................................................................... 366
14.4 Biogenic Fluxes in the Water Column.................................................................................................. 368
14.5 Bacteria .................................................................................................................................................. 371
14.6 Viruses ................................................................................................................................................... 376
14.7 Protozoa ................................................................................................................................................. 376
14.8 Bacteria–Protozoa–POM Interactions ................................................................................................... 382
14.9 Interactions of Bactivorous Gazers and Heterotrophic Bacteria .......................................................... 385
14.10 The Microbial Loop............................................................................................................................... 386
14.11 Nutrient Cycling .................................................................................................................................... 392

Chapter 15 Ecosystem Dynamics

15.1 Introduction............................................................................................................................................ 398
15.2 Pelagic Zonation .................................................................................................................................... 399
15.3 Phytoplankton and Primary Production ................................................................................................ 401
15.4 Food Webs ............................................................................................................................................. 406
15.5 Ecosystem Models ................................................................................................................................. 420
15.6 Southern Ocean Environmental Variability and Its Impact on the Pelagic Ecosystem....................... 442

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Chapter 16 Resource Exploitation
16.1 Introduction.............................................................................................................................................. 449
16.2 Krill .......................................................................................................................................................... 450
16.3 Fish........................................................................................................................................................... 454
16.4 Seals ......................................................................................................................................................... 458
16.5 Whales...................................................................................................................................................... 461

Chapter 17 Ecosystem Changes Resulting from Resource Exploitation

17.1 Introduction.............................................................................................................................................. 467
17.2 Ecosystem Changes Following the Decline in Whale and Fish Stocks................................................. 467
17.3 Changes in Seal Population Dynamics ................................................................................................... 470
17.4 Changes in Breeding Success in Birds.................................................................................................... 472
17.5 The “Whale Reduction–Krill Surplus” Hypothesis ................................................................................ 472
17.6 Potential Ecosystem Changes That Might Result from Future Resource Exploitation......................... 473

Chapter 18 Management of Living Resources

18.1 Introduction.............................................................................................................................................. 475
18.2 The Convention on the Conservation of Antarctic Marine Living Resources (CCAMLR).................. 475
18.3 Ecosystem Approach to the Management of the Living Resources of the Southern Ocean................. 476
18.4 The Role of Modelling Studies ............................................................................................................... 480
18.5 Monitoring Indicators of Possible Ecological Changes in the Antarctic Marine Ecosystem................ 483
18.6 Experimental Fishing as a Management Tool ........................................................................................ 486
18.7 Alternative Management Strategies ........................................................................................................ 486

Chapter 19 Ultraviolet Radiation

19.1 Introduction.............................................................................................................................................. 489
19.2 Biological Hazards of Ultraviolet Radiation........................................................................................... 490
19.3 UV Tolerance Mechanisms ..................................................................................................................... 491
19.4 Impacts of Ultraviolet Radiation on Antarctic Marine Biota................................................................. 492
19.5 Conclusions.............................................................................................................................................. 498

Chapter 20 Global Warming and Antarctic Marine Ecosystems

20.1 Introduction.............................................................................................................................................. 499
20.2 The Physical Environment ...................................................................................................................... 500
20.3 Evidence of Global Warming in the Southern Ocean ............................................................................ 502
20.4 The Potential Impact of Global Warming on Antarctic Marine Ecosystems ........................................ 509
20.5 Potential Changes in Ecosystem Structure and Dynamics ..................................................................... 516
20.6 Some General Conclusions...................................................................................................................... 517

Chapter 21 Human Impact

21.1 Introduction.............................................................................................................................................. 519
21.2 Impact of Waste Disposal ....................................................................................................................... 519
21.3 Impact of Tourism ................................................................................................................................... 521
21.4 Disturbance of Nesting Seabirds ............................................................................................................. 521
21.5 Contamination of Marine Biota by Pollutants ........................................................................................ 523
21.6 Impact of Oil Spills ................................................................................................................................. 524
21.7 The Impact of Resource Exploitation ..................................................................................................... 525

Epilogue ................................................................................................................................................................................... 527

Appendix.................................................................................................................................................................................. 531

References................................................................................................................................................................................ 533

Index......................................................................................................................................................................................... 609

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 The Southern Ocean
1
CONTENTS

1.1 Introduction ..........................................................................................................................................................................1

1.2 The Evolution of the Southern Ocean .................................................................................................................................3
1.3 Bathymetry ...........................................................................................................................................................................4
1.4 Climate .................................................................................................................................................................................4
1.4.1 Wind .........................................................................................................................................................................4
1.4.2 Temperature..............................................................................................................................................................4
1.4.3 Solar Radiation .........................................................................................................................................................5
1.5 Ice Cover ..............................................................................................................................................................................6
1.5.1 Seasonal Variation....................................................................................................................................................6
1.5.2 Ice Shelves and Icebergs..........................................................................................................................................6
1.6 Circulation Patterns and Water Masses ...............................................................................................................................7
1.6.1 Introduction ..............................................................................................................................................................7
1.6.2 Driving Forces ..........................................................................................................................................................8
1.6.2.1 Wind ..........................................................................................................................................................8
1.6.2.2 Ocean-Atmosphere Heat and Fresh Water Flux.......................................................................................9
1.6.3 Oceanic Circulation................................................................................................................................................10
1.6.3.1 Antarctic Circumpolar Current (West Wind Drift) ................................................................................10
1.6.3.2 Antarctic Coastal Current (East Wind Drift)..........................................................................................10
1.6.3.3 Circumpolar Frontal Zones .....................................................................................................................10
1.6.3.4 Gyres, Eddies, and Rings ........................................................................................................................10
1.6.3.5 Polynas.....................................................................................................................................................10
1.6.4 Water Masses .........................................................................................................................................................12
1.7 Some Regional Hydrographic Features .............................................................................................................................13
1.7.1 Southwest Atlantic and the Antarctic Peninsula Region.......................................................................................13
1.7.2 The Ross Sea ..........................................................................................................................................................13
1.8 Bottom Water Formation ...................................................................................................................................................14
1.9 Nutrients .............................................................................................................................................................................15

1.1 INTRODUCTION of the Southern Ocean (El-Sayed 1977, 1981, 1996;

Over the past two decades in particular there has been an
increasing emphasis on integrated studies of the Southern
Ocean (Figure 1.1) aimed at understanding what has been
termed “the Antarctic or Southern Ocean ecosystem.” While
the Southern Ocean can be considered a single ecosystem, it is
actually a series of interconnected ecosystems. These will be
discussed in subsequent chapters. Descriptions of the
Southern Ocean system have been given by several workers
over the years (Hart 1942; Currie 1964; Holdgate 1967; Knox
1970, 1983; Everson 1977a, 1984c; Bengtson 1978, 1985a;
Baker 1979; Tranter 1982; Hempel 1985a, 1985b, 1987)
The living resources of the Southern Ocean and their past
and future exploitation have been reviewed by the
SCAR/SCOR Group of Specialists on the Living Resources
Sahrhage 1988a), and they have also been the subject of
numerous reviews: Everson (1977b), Bengston (1978,
1985a), Knox (1983, 1984), and Anonymous (USSR)
(1984a, 1984b). The physical structure of the system has
been described by Deacon (1937, 1982, 1984b), Brodie
(1965), Gordon (1967, 1983), Gordon and Goldberg (1970),
Gordon et al. (1978), Forster (1981, 1984), Gordon and
Molinelli (1982), Amos (1984), Gordon and Owens (1987),
Squire (1987), Foldvik and Gammelsrad (1988), and Smith
(1990b). Environmental data for the Southern Ocean is
available from a variety of sources. Some of the most
valuable are the Discovery Reports, the Soviet Atlas of
Antarctica (Makismov 1966), the U.S. Navy Hydrographic
Office Oceanographic Atlas of the Polar Seas, the U.S.
National Center for Atmospheric Research Climate of the

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 2 Biology of the Southern Ocean

90°W
Ice edge (March)
West wind drift South America
East wind drift
Weddell sea drift Falkland Is.

Drake passage
60°W
e
nc
e rg e
rctic c onv South Shetland Is.
Anta South Georgia
ce
g en
er
div South Orkney Is.
ic
rct
ta
An South Sandwich Is.
Weddell sea

Filchner
60° Ice Shelf 60°
Ross Sea Ross 0°
180°
Antipodes Is. Ice Shelf
Bouvet Is.

Campbell Is.
Auckland Is. Antarctic continent
Wilkes
Macquarie Is. Land

Tasman
Sea

Tasmania Prince Edward Is.

60° Marion Is.

Heard Is. Crozet Is.

Australia

Kerguelen Is.

90° E

FIGURE 1.1 Antarctica and the Southern Ocean showing the positions of oceanic fronts and surface currents. (From Knox, G.A., Antarctic
Resources Policy: Scientific Legal and Political Issues, Vicuna, F.O., Ed., Cambridge University Press, Cambridge, 21, 1983. With
permission.)

Upper Air: Southern Hemisphere (Taljaard et al. 1971), and
the American Geophysical Union Antarctic Research Series
(1964a). The interlinking of the biological and physical
components of the system have been discussed by Knox
(1960, 1970, 1983, 1994), Holdgate (1967), Lubimova,
Naumov, and Lagunov (1973), Lubimova et al. (1980),
Deacon (1982), Lubimova (1982, 1983), Tranter (1982),
and Hempel (1985a, 1985b).
Hedgpeth (1977b) Points out that the Southern Ocean is
“a rich, apparently high productive plankton–pelagic
system supporting (at least in the past) great populations
of whales and millions of penguins, and seals, and
abundant intermediate populations of fish and cephalopods,
depending on the near surface productivity.” Among the
important characteristics of the Southern Ocean system are
the following:
1. It is a large system, indeed probably the largest
marine ecosystem on the globe.
2. It is semi-enclosed, especially in the overlying
water masses, and the Polar Front forms a distinct
northern boundary.

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 The Southern Ocean 3

3. It is an old system with a long evolutionary history
(Knox and Lowry 1977). The main circulation
patterns and water mass distributions were estab-
lished at least 20 million years ago (Knox 1980).
4. Most of the major taxonomic groups are circum-
polar in distribution. The principal variation is that
of productivity, which is greater in certain regions
than in others.
5. The quantitative and qualitative features of the
basic processes in the Southern Ocean system
differ obviously from those of other oceanic
systems, as demonstrated by the distribution of
the dominant herbivore and key species of the
system, Euphausia superba.
1.2 THE EVOLUTION OF THE SOUTHERN
OCEAN
The unique characteristics of the Southern Ocean are the
result of a long evolutionary history that can only be
understood within a palaeo-geographic framework of conti-
nental drift, plate tectonics and polar wandering (Knox 1979,
1980; Norton and Slater 1979; Kennett 1983; Elliot 1985;
Anderson 1999). Kennett (1977) has provided a detailed
synthesis of the information concerning the evolution and
palaeooceanography of the Southern Ocean, while Kemp
(1972) has discussed the broad trends in climate since
the Paleocene.
A summary of the major events in this evolution
follows: during the Cretaceous period equatorial seas
extended almost uninterrupted throughout the globe
(Frakes and Kemp 1972). In the Southern Hemisphere,
Antarctica (then part of Gondwanaland) was very close to
its present position near the rotational pole, but with
South America, New Zealand, and Australia attached
(Figure 1.2). New Zealand separated from Australia and
Antarctica in the Cretaceous, 60–80 million years before
present (my BP). By the late Paleocene (50 my BP) the
Tasman Sea had formed and Australia had begun to
separate from Antarctica. However, the South Tasman
Rise, which is of continental origin, was still part of

20°N Africa
South America
South America Africa
0°
0°
India
India

Antarctica Antarctica
lia
tra

Edge of continental shelf

a
ali
s

Mid-ocean ridge and associated faults

Au

South pole str

Au
(a) 20°S 40°S 60°S 80°S 40°S (b) 0° 20°S 40°S 60°S 60°S 40°S 20°S

Africa Africa
South
0° India South
America
America

Antarctica
Antarctica
a
ali
a

str
ali

Au
str
Au

(c) 0° 20°S 40°S 60°S 60°S 40°S 20°S (d) 0° 20°S 40°S 60°S 60°S 40°S 20°S

FIGURE 1.2 Stages in the dispersal of the Gondwana continents (after Norton and Sclater 1979). No account is taken of the possibility of
rearrangement of West Antarctica. Note that only a very limited separation of Australia and Antarctica had occurred by 53 my BP, and also
that South America and the Antarctic Peninsula were most likely still abutting each other at 39 my BP. The position of the South Pole is that
inferred for the time given in each diagram. (From Elliot, D.H., Antarctica, Key Environment Series, Pergamon Press, Oxford, 39, 1985.
With permission.)

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 4 Biology of the Southern Ocean
Victoria Land, and because of this no major current
system could develop between Australia and Antarctica.
Frakes and Kemp (1972) postulated that during the
Eocene large oceanic gyres extended from the equator
to high latitudes and brought warm water to the Antarctic
coastline. Their reconstructed palaeotemperatures at
608S latitude are 248S for Queen Maude Coast, 178C
for the shallow sea between Australia and Antarctica,
and 78C for the attached South American–Antarctic
Peninsula coastline.
In the Late Eocene-Early Oligocene (39 my BP),
major changes began to occur which transformed the
global warm climate into the modern climatic regime of
today. At that time there was a dramatic lowering
of austral temperatures. This resulted in near-freezing
surface coastal waters along the Antarctic coastline,
which probably caused sea level glaciation around the
continent, and the formation of cold Antarctic Bottom
Water. Associated with the development of the thermo-
clinal circulation system was an increase in the calcium
carbonate compensation depth (CCD) (Heath 1969; Van
Andel and Moore 1974). Evidence from sediment macro-
fossils (foraminiferans, diatoms, and radiolarians)
indicated a marked change to a cold water biota (Margolis
and Kennent 1970; Jenkins 1974).
In the Late Oligocene (25–28 BP) the South Tasman Rise
finally separated from Victoria Land sufficient enough to
allow the formation of the Antarctic Circumpolar Current
(West Wind Drift) (Figure 1.1). The Drake Passage had
formed some time between the initial separation of Australia
from Antarctica and the final separation of the Tasman Rise.
In the Early Miocene (22 my BP) the Polar Front (Antarctic
Convergence) formed, producing a major biological barrier
which is still operating today. Large ice sheets developed
rapidly in East Antarctica during the Middle Miocene. Dell
and Fleming (1975), using fossil molluscan evidence,
suggested that the sheltered rocky coastline of the Ross Sea
was, at that time, probably ice-free and kelp-fringed, not
unlike Tierra del Fuego today. By the Late Miocene-Early
Pliocene, the West Antarctic ice sheet had formed and was
much thicker than it is today (Shackleton and Kennett 1975).
During this time the production of siliceous phytoplankton
steadily increased. Hays (1969) observed two intervals of
distinct cooling at 2.5 and 0.7 my BP, and climatic oscil-
lations during the last 3.5 million years, with a steadily
increasing coolness up to the present time.
1.3 BATHYMETRY
The bathymetry of the Southern Ocean is shown in
Figure 1.3. Three deep-water basins (4,000–6,000 m deep)
surround the Antarctic continent: the Atlantic–Indian Basin,
the Indian–Antarctic Basin, and the Pacific–Antarctic Basin.
These basins are partially bounded on the north by a series of
ridges or rises; the Scotia Ridge and the Atlantic–Indian
Ridge, the Southeast Indian Ridge, and the Pacific Atlantic
Ridge respectively. These ridges and the Kerguelen Plateau
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tend to restrict the flow of bottom water and in some areas
they even deflect surface currents. The Drake Passage
between South America and the Antarctic Peninsula restricts
the circulation of water masses and, as will be discussed
later, has a profound effect on circulation in the
Southern Ocean.
The continental shelf of the Southern Ocean differs
from that surrounding other continents in that it is
unusually deep, with the “shelf break” (the transition
between the continental shelf and the continental slope)
lying two to four times deeper than in other oceanic
regions. This is partly due to the isotactic equilibrium
adjustment of the continent to the large mass of the
Antarctic ice sheets. Both the Weddell Sea and the Ross
Sea are characterized by broad ice shelves: the Filchner
and Ronne Ice Shelves and the Ross Ice Shelf respect-
ively. These shelves, as we shall see, profoundly influence
the near-shore circulation and water properties.
1.4 CLIMATE
1.4.1 WIND
A ring of low pressure surrounds the Antarctic continental
plateau, while tropical anti-cyclones lie to the north.
Winds in the Southern Ocean blow towards the trough
of low pressure, but are directed to the left by the earth’s
rotation (Coriolis effect). As a result, at about 508S of the
Southern Ocean, westerly winds occurs. The westerlies
within this circumpolar belt are quite strong, with
maximum intensity in the region of the Antarctic Circum-
polar Current (ACC) (see Figure 1.6). It is this wind that
drives the Southern Ocean circulation. Superimposed upon
this westerly circulation are northwest–southeast
moving depressions.
Over the Antarctic continent itself, katabatic winds
dominate the weather system. These winds, driven by very
cold and dense air flowing down the glaciers and ice streams
of the ice cap, can often reach very high velocities, and they
are also responsible for the strong southeast–northwest winds
extending many kilometers from the coast out to sea. Near
the peri-Antarctic trough of low pressure there is a belt of
easterly winds.
To summarize, the prevailing wind system of the
Southern Ocean comprises three main elements: a south-
easterly component near the coast, a zone of easterly flow
encircling the continent and extending north to about 658S,
and a wide zone of westerlies reaching as far north as about
408S (Squire 1987). This zonal circulation pattern is more
intense and constant here than in any other region of
the globe.
1.4.2 TEMPERATURE
The Southern Ocean is cold. Across the Polar Frontal Zone
the temperature range in the summer is from 4 to 88C, and in
 The Southern Ocean 5

Atlantic
Indian Basin
Argentine
Basin Atlantic Indian

Ridge

Indian Crozet
Weddell Antarctic Basin
Basin Basin

n
le
g ue au
r t e
Ke Pla

ian
Ind
Pacific
Antarctic

ge
st
ea
Basin

Rid
uth
So
Indian
Pacific Antarct Antarctic
ic R
id Basin
ge

Australian
Basin
Southwest
Pacific
Basin

FIGURE 1.3 Map of the sea bed surrounding Antarctica showing the principal deep water basins and submarine ridges. (From Squire, V.A.,
Primer Symposium Espanol de Estudiios Antarcticos Palma de Mallorca June–July, 201, 1987. With permission.)

the winter from 1 to 38C. Surface waters south of the Polar
Front have an average temperature of about 1–28C in the
winter and 3–58C in the summer, while further south near the
continent temperatures range from only about K1.0 to
K1.98C. Temperature differences between cold surface and
bottom layers and intermediate warmer layers is less than
58C; thus the total annual range throughout does not exceed
4–58C and is considerably less for the greater part of the area
(e.g. in McMurdo Sound temperatures range seasonally only
from K1.7 to K1.98C).
1.4.3 SOLAR RADIATION
In the far south near the continent, the alternation between
total darkness for half of the year and continuous daylight for
the other half imposes a seasonal light regime in contrast to
the diurnal cycles of lower latitudes. In addition, the amount
of light penetrating the surface waters of the ocean is
determined not only by its intensity, angle of incidence,
surface reflection (up to 50% according to data given in 1967
by El-Sayed for Marguerite Bay in February 1965), and

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 6 Biology of the Southern Ocean
absorption by suspended particles, but also by the presence of
sea ice and snow cover (see Section 2.8.7). Light penetration
is also influenced by the transparency of the atmosphere.
Anti-cyclonic conditions near the coast result in skies that are
often lightly cloudy or clear. In the region of the ACC there is
a continuous passage of low pressure systems and a predo-
minance of cloudy weather resulting in lower insolation in
this region relative to areas close to the continent (Holm-
Hansen et al. 1977).
Oceanic waters surrounding Antarctica are typically blue
and highly transparent with a maximum Secchi depth (the
depth at which a standard Secchi disc becomes invisible) of
about 40 m (Slawyk 1979). The 1% light level is relatively
deep at about 100 m. Tilzer et al. (1986) attribute the low
concentrations of dissolved organic compounds and the low
abiotic turbidity to the very low terrestrial input of sediment
and organic matter.
1.5 ICE COVER
1.5.1 SEASONAL VARIATION
One of the salient characteristics of the Southern Ocean is the
dramatic changes that occur in sea ice cover, from about
20!106 km2 in late winter to about 4!106 km2 in late
summer (Zwally et al. 1983a,1983b; Comiso and Zwally
1984). The ways in which sea ice forms and decays and the
characteristics of the different types of sea ice are considered
in Section 3.2. While the general distribution of sea ice has
been known for many years from scattered ship observations,
it is only recently that continuous year-round observations
have been available as a result of satellite imagery, thus
enabling the detailed changes over the year and year-to-year
variations to be documented.
Figure 1.4 illustrates the growth and decay of the sea ice
in the Southern Ocean. The months of minimum extent
are February and March, while those of maximum extent
are September and October. The northern limit of the sea ice
changes by no more than a few degrees from year to year.
The most rapid advance of the ice edge occurs in May and
June, when it moves northward at a rate of 4.2 million km2
per month; November and December are the months of the
most rapid retreat when the ice edge recedes at
6.9 million km2 per month (Squire 1987). The average
rates of advance and retreat are respectively 2.4 and
3.3 million km2 per month. Figure 1.5 depicts the average
seasonal cycle of the monthly Southern Ocean sea ice over
the period 1978–1987, which was calculated from Nimbus 7
satellite imagery.
Over time the sea ice records show considerable
variability from year to year in: (1) sea ice extent, (2)
the spatial range experienced in the monthly average
Southern Ocean sea ice distribution, and (3) the length
of the sea ice season. Using the Nimbus 7 data, Parkinson
(1992) has examined this variability. Regarding the three
variables listed above, (1) maximum sea ice extent varied
by approximately 12%, decreasing during the mid-1970s,
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followed by increases over the next few years and
leveling off for much of the 1980s, and (2) the area of
intra-annual variability in monthly average sea ice distri-
butions in summer far exceeded the summer-time area of
consistent ice coverage, this in sharp contrast to the
winter-time situation, when the area of consistent sea
ice coverage is considerably larger. In winter, the sea
ice distribution variability is largely confined to two
regions: a relatively narrow band, generally 2–58 of
latitude, surrounding the region of consistent ice coverage,
and, for the mid-1970s, the region of an occasional large
open water area within the pack ice in the Weddell Sea,
termed the Weddell polyna. The length of the sea ice
season, calculated for the years 1979–1986, with satellite
passive-microwave data coverage through all months of
the year, showed increases over the period in the Ross
Sea but decreases in the Weddell and Bellinghausen Seas.
In both cases it appears through comparisons with data
from 1973–1976, that the 1979–1986 changes more likely
reflect a fluctuating behavior of the ice cover than a
long-term trend.
At the minimum extent scattered areas along the
Antarctic coast retain some ice coverage. Most of this ice
is found in the eastern Weddell Sea and the Bellinghausen–
Amundsen Sea sector. No ice is found north of the Ross Ice
Shelf at this time. Around the margins of the continent areas
of unbroken fast ice (ice attached to the shore) may persist for
two or more years, forming thick multi-year ice. This is in
contrast to the pack ice (drifting ice floes) zone of annual sea
ice, which is 1–2 m in thickness. The distribution of ice in the
pack ice zone is highly variable as the ice cover is frequently
broken up by storms, and the resulting floes, driven by
surface currents and wind, drift considerable distances.
Leads of open water (open channels in the ice within the
pack) are highly variable.
1.5.2 ICE SHELVES AND ICEBERGS
The Antarctic continent is largely covered by ice, and
around the margins the extensive ice sheets extrude from
the continent into the sea. In the Ross and Weddell Seas
these ice sheets float on the ocean forming the extensive
Ross, Filchner, and Ronne Ice Shelves. The layer of water
under the Ross Ice Shelf varies from a few meters to
several hundred meters thick. The ice layer varies from
200 m thick at the seaward edge to over 800 m at the
grounded edge. The ice shelves are continually moving
seaward at about 1 m per day due to the accumulation of
snow on the continent. As they extend into the open
ocean, they are exposed to the action of long period
waves and eventually they crack and calve icebergs. The
large tabular icebergs so produced can range from a few
hundred meters up to 100 km in horizontal extent and
usually are about 200–300 m thick. It has been estimated
(Radok, Streten, and Weller 1975) that the total mass of
icebergs is about one-third of the mass of sea ice at
 The Southern Ocean 7

90°

60° 120°

30° 150°

0° 50° 60° 70° 80° 85° 180°

30° 150°

March September
60° 120°
1974

90°

FIGURE 1.4 The distribution of the pack-ice in winter (September) and summer (March). (From Knox, G.A., Ocean Management, 9, 113,
1984. With permission.)

maximum extent. The average life of icebergs is about
four years, but large ones have been tracked for much
longer periods (Swithinbank, McClain, and Little 1977).
Figure 1.6. gives estimates of the variation of Antarctic
ice sheet parameters with time. It can be seen that a rapid
change in the mass budget of the Antarctic ice sheet
occurred around 10 my BP when the total budget was two
to three times its present level (Robin 1988). The curve
“total discharge to the sea” is ice lost by basal melting
beneath the ice shelf and ice tongues and by the calving of
icebergs. Both of these have decreased markedly since that
period. The extent of the pack ice (Figure 1.6c) shows the
large increase in extent, especially during winter, since the
opening of the Drake Passage resulted in Antarctic waters
becoming thermally isolated as the circumpolar current
developed (Robin 1988).
1.6 CIRCULATION PATTERNS AND WATER
MASSES
1.6.1 INTRODUCTION
The basic feature of the circulation patterns and hydro-
graphic processes in the Southern Ocean were first
described by Deacon (1937, 1963, 1964a, 1964b, 1977,
1984b). Since then there have been a number of reviews
including those of Gordon, George, and Taylor (1977a,
1977b), Forster (1981, 1984), Gordon and Molinelli

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 8 Biology of the Southern Ocean

2.0 below the main thermocline of the world’s oceans. Associ-

ated with this is significant heat flux across the ACC.
While the summary of circulation patterns given below
1.5 represents a generalized picture of the system dynamics, it
Ice extent (106 km2)

should be emphasized that, superimposed upon this, is a great

1.0
5 complicates water mass properties in two ways: (1) the
0 in regard to momentum, heat, water and gas exchange, and
J F M A M J
FIGURE 1.5 Average seasonal cycle of monthly average Southern
Ocean sea ice extents over the period with full-month coverage of
Nimbus 7 SMMR. November 1978 through July 1987. Ice extents
are calculated from passive-microwave SMMR data as the aerial
coverage of the Southern Ocean with ice concentrations exceeding
15%. (From Wadhams, P., Ice in the Oceans, Gordon & Breach
Science Publishers, London, 45, 1967. With permission.)
(1982), Gordon (1983, 1988), Amos (1984), Gordon and
Owens (1987), Squire (1987), and Carmack (1992).
The Southern Ocean-atmosphere-ice coupled system is
extremely complex. Processes within the Southern Ocean
are responsible for the production of water characteristics
J A
amount of temporal, seasonal, and year-to-year variability
(see Chapter 15). The Southern Ocean is not radically
symmetric, and many of the circulation and water mass
features vary markedly with longitude. The cryosphere
highly spatially and temporarily variable sea ice cover
strongly influences the coupling of ocean and atmosphere
S O N D (2) ocean interaction with glacial ice has a marked influence
on the characteristics of water masses (Gordon 1988).
1.6.2 DRIVING FORCES
1.6.2.1 Wind
In general, the principle driving force for the Southern
Ocean circulation is the wind field. The westerlies within
the circumpolar belt are quite strong with the maximum
westerlies situated in close proximity to the ACC. The
wind field produces Ekman divergence (upwelling) south
of the ACC and convergence (sinking) to the north.
The upwelling poleward of the ACC carries about 45 Sv
(1SvZ1!106 m2 s1) into the mixed layer, two-thirds of

Ice volume (km3 × 107) Glacier ice budget (km3 × 103) Pack ice (km2 × 106)
0 1 2 3 4 0 1 2 3 4 5 0 10 20 30
0 0 0

Summer
20 20 20 Winter
Total discharge
to sea
Glacier berg discharge
40 40 40
Million years BP

Million years BP

Million years BP

Total mass budget

60 60 60

80 80 80

100 100 100

(a) (b) (c)

FIGURE 1.6 Estimated variation of the Antarctic ice sheet parameters with time. (a) Volume of ice sheet including the ice shelves; (b) Annual
(total) mass budget of the Antarctic ice sheet, together with annual ice discharge to sea across the flotation line, and the annual volume
discharged by glaciers directly to open water but not to ice shelves (glacier berg discharge); (c) Changes of the area extent of pack ice around
Antarctica. (From Robin, G. DeQ., Palaeogr. Palaeoclimat. Palaeoecol., 67, 45, 1988. With permission.)

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 The Southern Ocean 9

which is directed the north, with the rest towards the
continent, where coastal sinking occurs (Gordon 1988).
Along the margins of the continent, south of about 608S,
the wind field results in westerly flowing water—the
Antarctic Coastal Current (East Wind Drift). Between the
two well-defined current systems there is a series of eddies.
Figure 1.7 shows the general pattern of surface currents in the
Southern Ocean. Since the water column is weakly stratified
south of the Polar Front, the ocean currents extending to the
bottom result in the deep circulation being similar to that of
the surface circulation, although the former is more strongly
influenced by bottom topography.
1.6.2.2 Ocean-Atmosphere Heat and Fresh
Water Flux
Within the region of the Southern Ocean there is a slight
excess of precipitation over evaporation (Gordon 1981). The
annual freezing and thawing of the sea ice is a significant
factor in the redistribution of freshwater. In addition,

°E
30°

30
W

Antarcti c circu
mpola
r cu
rre
nt °E
Antarctic 60
divergence
16
0°
W
Antarctic co
a s ta
lc
Weddell
ur
re
sea
nt

gyre

90° W 90° E

Ross
sea
gyre

12
W 0°
0° E
12
180°
Antarctic
convergence
Subtropical
convergence
15
0°
E
W 0°
15

Warm currents
Cold currents

FIGURE 1.7 Surface currents in the Southern Ocean and the mean positions of the principal frontal zones. Warm currents are shown dotted
and cold are shown solid. (From Knox, G.A., Antarctic Resources Policy: Scientific, Legal and Political Issues, Cambridge University Press,
Cambridge, 21, 1983. After Brodie [1975], adapted from Sverdrup, Johnson, and Fleming [1942]. With permission.)

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 10
the waters of the Southern Ocean play a primary role in the
wastage of glacial ice, mainly by direct melting of the ice
along the underside of the shelf ice (Jacobs, Fairbanks, and
Horibe 1985; Schlosser 1986).
As Gordon (1988) points out, estimates of ocean-
atmosphere flux are subject to large errors due to the
lack of meteorological and sea ice (concentration, thick-
ness) data, and especially the paucity of data from the
winter period. The strong seasonal variability of the sea ice
cover, coupled with approximately 10–20% inter-annual
variability in maximum winter sea ice cover (Zwally et al.
1983a, 1983b), further complicates estimates of the mean
ocean-atmosphere energy exchange. South of 608S, the
northern limit of sea ice cover, heat loss is large an
annual average of 30 W m2 (Gordon 1981) but it is
highly dependent on ice cover. Heat flux across the ACC
and Polar Front (average 558S) is 3.1!104 W, while that
across 608S is larger, amounting to 5.4!104 W (Gordon
and Owens 1987). In order to maintain a steady state
condition, the net poleward heat flux must involve the
poleward transport of a large volume of water.
1.6.3 OCEANIC CIRCULATION (FIGURE 1.7)
1.6.3.1 Antarctic Circumpolar Current (West
Wind Drift)
The Antarctic Circumpolar Current (ACC) is unique in that it
is the only zonal current encircling the globe, and it is almost
unobstructed by land masses, being constricted only in the
region of the Drake Passage. It displays significant variations
with longitude and is principally related to bottom topo-
graphy, being deflected where it passes over submarine
ridges with a strong northerly component after it passes
through the Drake Passage. Its width varies from less than
200 km south of Australia to over 1,000 km in the Atlantic.
Mean current speeds within the ACC are relatively low
(0.04–0.25 m sK1 , some 2–3% of the wind velocity).
Although the predominant movement is to the east, there is
a strong northerly component. There is also some evidence
that there are meanders and even loops within the current
(Tchernia 1974, 1980; Joyce and Patterson 1977). The ACC
joins the great current gyres of the South Atlantic, South
Indian, and South Pacific Oceans.
1.6.3.2 Antarctic Coastal Current (East Wind
Drift)
In the vicinity of the Antarctic continent south of about 658S,
where winds from the east and southeast blow off the ice
sheet, there is a surface current which flows westward. This
Antarctic Coastal Current generally follows the coastline
with two important indentations in the Ross and Weddell
Seas. Its flow is estimated at 10 million m3 sK1, while its
speed is variable between 0.1 and 1 m sK1, with the largest
values near the Ross Sea Shelf (Squire 1987). The current
may not be continuous but rather broken up by a series of
gyres (Figure 1.7).
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Biology of the Southern Ocean
1.6.3.3 Circumpolar Frontal Zones
There are number of fronts within the Southern Ocean
(Deacon 1982). While some are circumpolar, such as the
Polar Front (Antarctic Convergence) they display much
variability in characteristics with longitude. The most
significant of these fronts is the Polar Front which
divides the Southern Ocean into two distinct regions:
SubAntarctic to the north and Antarctic to the south. It is
a zone of variable width characterized by steep gradients in
sea-surface temperature, abrupt changes in phytoplankton
abundance, zooplankton distribution, pelagic bird species,
weather conditions, and sometimes by a salinity maximum
at the surface. A circumpolar subsurface salinity minimum
closely follows the surface position of the front (Gordon,
George, and Taylor 1977a, 1977b; Joyce, Zenk, and Toole
1978; Sciermammano 1989).
To the north of the Polar Front lies the SubAntarctic
Front, while to the south lies the Antarctic Divergence. This
divergence is caused partly by the northward component
associated with the ACC and the southward component of
the Antarctic Coastal Current, and partly by circulatory and
thermohaline factors of the southern density gradient and the
distribution of pressure and winds. In the region of the
divergence deep water upwells to reach the surface.
The classical schematic diagram of the horizontal and
vertical movements of the Antarctic circumpolar water is
presented in Figure 1.8.
1.6.3.4 Gyres, Eddies, and Rings
A characteristic of the zonation of circumpolar currents is the
development of eddies of variable size and duration. This
eddy variability is due to a combination of factors (Gordon
1988). The current cores migrate laterally (Nowlin, Whit-
worth, and Pillsbury 1977) by as much as 100 km in 10 days.
Meanders or waves form and propagate along the ACC fronts
(Legechis 1977; Sciermammano 1989). These meanders
sometimes develop into closed rings (Joyce and Patterson
1977). Cold and warm water core rings have both been
observed. The eddies are generally 30–100 km wide with
surface velocities typically 30 cm sK1 or more, and they are
vertically coherent from the surface to the bottom. Numerous
current rings and meanders have been reported from the
Drake Passage.
In addition to eddies there are large permanent cyclonic
flowing gyres. The largest and best-defined of these is the
Weddell Gyre, extending east of the Antarctic Peninsula to
about 208W and from Antarctica near 70–608S (Deacon 1979;
Gordon, Martinson, and Taylor 1981; Comiso and Gordon
1987). Others are the Ross Sea Gyre, north and east in the Ross
Sea (Tchernia and Jeannin 1984; Gouretski 1999). A review of
the Weddell Gyre system is found in Deacon (1977).
1.6.3.5 Polynas
As the sea ice cover expands in winter, areas of open water
occur polewards of the ice limit which do not freeze but
 The Southern Ocean 11

70°W 60 50 40 30 20
35°S
00

00
30

10
00 SACW
50
STF

Brazil cu
40

r r ent

50
00
Falklan d current

p
Argentine Basin

ga
45

ge
5000

rid
d
an
lkl
SAF

Fa
SAZ
Falk
l and Ridge
50 escarpm
en te Falkland PF
Ewing
Falkland Bank Basin

Isl
Falkland

as
ia
(Malvinas) plateau Georg

orc
islands

ad
Scotia Ridge
r th

as
Burdwo
od No

ris
bank South

e
55 Georgia
Scotia sea
PFZ

SAF
AZ
Drake passage
60

FIGURE 1.8 The upper ocean water masses and fronts in the western South Atlantic, SACW, South Atlantic Central Water; PF, Polar Front;
SAZ, Subantarctic Zone; STF, Subtropical Front; PFZ, Polar Frontal Zone; SWAF, Subantarctic Front; AZ, Antarctic Zone; The broken line
around the Ewing Bank is the 2,000 m isobath. (From Stein, M., Heywood, R.B., Southern Ocean Ecology: The BIOMASS Perspective,
Cambridge University Press, Cambridge, 12, 1996. With permission.)

instead remain open to the atmosphere through all or part of
the winter. These regions of long-lived open water are called
polynas. Polynas tend to occur where high winds rapidly
disperse freshly-formed ice at times when air temperatures
are well below the freezing point of sea water (Van Woert
1999). Polynas play an important role in heat transfer from
the oceans to the atmosphere, ice production, the formation
of dense shelf water, spring disintegration of sea ice,
phytoplankton and zooplankton production, and the distri-
bution of higher trophic animals such as cephalopods, fish,
birds, seals, and cetaceans. Within polynas, the oceanic heat
loss may be 10–100 times above that of the ice-
covered surface.
A review has categorized two distinct types of polynas
(Smith et al. 1990b). A latent heat polyna is formed when sea
ice is continually removed from the region in which it is
formed by winds or ocean currents, the heat needed to
balance the loss to the atmosphere being provided by the
latent heat of fusion of the ice which continually forms.
A sensible heat polyna is formed when a continued source
of heat from the ocean prevents ice formation.
Coastal polynas are a common feature of the continental
shelf round the Antarctic continent. They are primarily latent
heat polynas where heat loss from the ocean surface is
balanced by the latent heat of new ice formation and the
polyna is maintained by wind or current removal of the new
ice. The coastal polynas around Antarctic can be further
subdivided into two categories: those off the front of major
bay ice shelves (the Ross, Filchner–Ronne, Amery etc.) and
those in the lee of ice tongues or other coastal projections such
as the Terra Nova Bay polyna in the Ross Sea (Bromwich, Liu,
and Rogers 1998) or along the Wilkes Land coast. Those off
the front of the ice shelves involve processes of water
circulation under the shelves and katabatic winds, whilst
inhibition of ice drift into polyna areas is important for
those in the lee of ice tongues (Smith et al. 1990).
Processes inside latent heat polynas are significant for the
sea ice zone as a whole. Being regions of intense heat loss
from the ocean to the atmosphere, these polynas can behave
as “ice factories,” contributing a significant fraction of the
total annual sea ice production; it has been estimated, for
instance that the small polyna in Terra Nova Bay contributes

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 12 Biology of the Southern Ocean

(°C)
rat ure 0
mpe Anta
face te rctic Ice
Sur 2 An Dive shelf
6 4 surf tarctic rgen
ce
8 ace
Anta wate
10 rctic r Shelf
con water
verg
Sub en ce z
0 surf antarct one
ace ic
wat
er
(Upper)
1000

slo p e
c e
r c ti a t
2000 S u b a n t a i at e w
i nte r m e d

en t a l
Depth (m)

te r

nti n
wa
3000 e ep

Co
ar d
um p ol
C i rc

te r
tic
(Lower)

rc
4000

wa
ta
An

m
tt o
Bo
80°
5000

FIGURE 1.9 Schematic diagram of the meridional and zonal flow and water masses of the Southern Ocean. The diagram represents average
summer conditions and applies particularly to the Atlantic sector. (From Knox, G.A., Antarctic Resources Policy: Scientific, Legal, and
Political Issues, Vicuna, F.O., Ed., Cambridge University Press, Cambridge, 113, 1983. After Brodie [1965], adapted from Sverdrup, Johnson,
and Fleming [1942]. With permission.)

10% of the ice production in the Ross Sea (Kurtz and
Bromwich 1985). Brine rejected during the ice growth is
concentrated in the polyna areas and can cause intense
localized water mass modification as well as significantly
increasing the salinity of the Antarctic shelf water. The ice-
free polynas also play an important role for Antarctic marine
ecosystems and in the control of biogeochemical fluxes.
A summary of recurrent polynas that have been detected
and studied using passive microwave data has been given by
Zwally, Comiso, and Gordon (1985). These include the
coastal polyna in the Cosmonaut Sea (Comiso and Gordon
1987, 1996) and in Terra Nova Bay in the Ross Sea (Kurtz
and Browich 1983; Bromwich, Liu, and Rogers 1998).
Zwally, Comiso, and Gordon (1985) and Bromwich, Liu,
and Rogers (1998) have reviewed the occurrence, extent, and
variability of the Ross Sea polyna, which covers an area of
about 27,000 km2. This polyna owes its existence to a
persistent cyclone near Roosevelt Island, which induces a
sea level pressure distribution over the Ross Ice Shelf. This
phenomenon results in an intensification and northward
propagation of katabatic winds across the ice shelf and out
to the sea. The immediate effect of such katabatic surge
events is the development of the Ross Sea Polyna.
Other polynas which also occur in the Ross Sea include
small polynas off Cape Royds, Ross Island, and a larger one
in Terra Nova Bay (Kurtz and Bromwich 1985; Van Woert
1999). The Terra Nova Bay polyna is a stable feature,
recurring annually and persisting throughout the winter,
although it appears to vary in extent from less than a
kilometer to 5,000 km2. Van Woert (1999) suggests that
the extent of the polyna is due to both local and katabatic
winds at Terra Nova Bay and regional katabatic winds
blowing off the Ross Ice Shelf.
1.6.4 WATER MASSES
Figure 1.9 presents an idealized schematic diagram of the
meridional and zonal flow in the Southern Ocean. There are
three principal hydrographic regimes: the area north of the
Polar Front, the area south of the Polar Front, and the area on
the continental shelf round Antarctica. Excluding the surface
waters, three water masses dominate the deep ocean: the
Subantarctic Intermediate Water north of the Polar Front
near the 1,000 m level, the Antarctic Bottom Water near the
bottom, and the Antarctic Circumpolar Deep Water in
between at various depths.
Warm Deep Water upwells at the Antarctic Divergence.
Here the oxygen minimum and the temperature maximum
coincide and rise together with the underlying salinity
maximum. The cold, low-salinity, high-oxygen Antarctic
surface waters move north to the Polar Front where they
sink and contribute to the formation of the low salinity
Intermediate Water. North of the Polar Front is the warmer
SubAntarctic Surface Water.

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 The Southern Ocean
Near the bottom of the ocean, very cold Antarctic Bottom
Water moves north from its primary site of generation around
the continental margin, especially in the Weddell Sea.
Sandwiched between the two Antarctic water masses is the
broad compensatory flow of high-salinity Warm Deep Water.
The south-flowing Deep Water mixes with the Antarctic
waters above and contributes to the Circumpolar Deep Water
which, in terms of volume, is the dominating water mass of
the Southern Ocean. Mixing takes place across the
boundaries between the water masses.
1.7 SOME REGIONAL HYDROGRAPHIC
FEATURES
1.7.1 SOUTHWEST ATLANTIC AND THE ANTARCTIC
PENINSULA REGION
Stein and Haywood (1994) have reviewed the physical
oceanography of the Southwest Atlantic region and the
Antarctic Peninsula area (Figure 1.8). On leaving the
confines of the Drake Passage and entering the Scotia
Sea the Antarctic Circumpolar Current (ACC) is deflected
north and west by the submerged ridge of the Scotia Arc,
before it turns east again to flow round South Georgia and
across the Atlantic Ocean sector of the Southern Ocean
(Deacon 1937; Gordon and Goldberg 1970). It consists of
Subantarctic and Antarctic Surface waters from the Pacific
Ocean sector of the Southern Ocean and from the Bellin-
ghausen Sea (Nowlin and Clifford 1977). Some mixing
with Weddell Sea Surface Water occurs in the Scotia Sea.
However, the southern arm of the Scotia Arc effectively
restricts the flow of water out of the Weddell Sea (Carmack
and Foster 1975) and most enters the Scotia Sea across the
Weddell–Scotia Confluence east of the South Orkney
Islands (Gordon 1967). Consequently there is a marked
gradient of mixing across the Scotia Sea with the influence
of the Weddell Sea Surface Water being greatest to the
south and east of South Georgia (Anonymous 1983;
Heywood, Everson, and Priddle 1985).
From north to south a number of fronts separate distinct
water masses. The most northerly front, the Subtropical
Front, marks the northern boundary of the sub-Antarctic
Zone, while the Subantarctic Front marks its southern
boundary. South of this front is the Polar Frontal Zone
with its southern boundary marked by the Polar Front. To
the south lies the Antarctic Zone. This complex of fronts and
water masses (Zones) is the result of the passage of water
through the Drake Passage being influenced by the various
banks and ridges: the Burwood Bank, the North Scotia
Ridge, the Ewing Bank, and the Falkland Ridge.
1.7.2 THE ROSS SEA (FIGURE 1.10)
Three main oceanic fronts occur within the Ross Sea region:
the Polar Front (Antarctic Convergence), the Antarctic
Divergence, and the Antarctic Slope Front. These fronts
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13
separate water masses of different temperature and salinity,
and they exhibit marked biological and physical changes.
The interchange of these water masses, especially at the
Antarctic Divergence, provides the minerals and nutrients
fundamental to marine biological production.
The Polar Frontal Zone lies between about 558S and 608S
and separates Subantarctic Water to the north from Antarctic
Water to the south. This is a zone of variable width,
characterized by steep gradients in sea surface temperature
and abrupt changes in phytoplankton composition,
zooplankton distribution, pelagic bird species, and weather
conditions. The Polar Front lies within the Antarctic Circum-
polar Current (West Wind Drift). It is a complex system
where wave-like disturbances frequently close on themselves
to form rings and eddies with lifetimes that vary from days to
years (Forster 1984).
South of about 658S, where winds blow from the east
and southeast off the ice sheet, the Antarctic Coastal
Current (East Wind Drift) flows in the opposite direc-
tion—from east to west. The boundary between the East
180° 170°W
Antarctic convergence 60°S
West wind drift
65°S
Antarctic divergence
East wind drift
70°S
Anta
rctic
Sl
op
ef
ro
nt 75°S
Ross
Ross ice shelf
FIGURE 1.10 Oceanic fronts and surface water circulation in the
Ross Sea region. Three main oceanic fronts occur: the Polar Front
(Antarctic Convergence), the Antarctic Divergence, and the
Antarctic Slope Front. (From Knox, G.A., Ross Sea Region: State
of the Environment Report for the Ross Sea Region of Antarctica,
Antarctic Institute, Christchurch, 54, 2001. With permission.)
 14
and West Wind Drifts forms the second frontal zone, the
Antarctic Divergence, where deeper water rich in nutrients
upwells to the surface, resulting in enhanced
phytoplankton production.
A third frontal zone, the Antarctic Slope Front, lies over
the edge of the continental shelf in the southern Ross Sea
(Ainley and Jacobs 1981; Jacobs 1991) and it is centered
10–45 km seaward of the shelf break, placing it just south of
the 1,000 m depth contour. This topographically controlled
front is marked by strong subsurface gradients in ocean
currents and temperature, and by along-shore currents that
are stronger than those occurring in most surface waters of
the adjacent continental shelf. The slope front lies at the
northern margin of the massive phytoplankton bloom that
occurs over the Ross Sea continental shelf in summer.
Warm deep water approaches the continental slope of the
Ross Sea from the northeast in the west and from the east-
northeast in the east. These waters are seasonally cooled and
warmed, Stalinized by the formation of sea ice, and fresh-
ened by water from melting ice. Ice and water is then moved
west and north by surface circulation, which is mainly due to
the extensive cyclonic gyre on the western side of the Ross
Sea. Surface waters move to the southeast along the eastern
periphery of this cyclonic system. As the southeast flowing
water approaches the Ross Ice Shelf, the current is deflected
to the west, with part joining the cyclonic circulation and part
passing beneath the shelf. Along the Victoria Land coast, a
northward flowing coastal current forms a well defined and
comparatively narrow stream (Tressler and Ommundsen
1962). Some of this water returns to the cyclonic circulation,
while the rest turns round Cape Adare and eventually joins
the Antarctic Coastal Current.
Subsurface circulation is formed by two different anti-
cyclonic gyres at each end of the Ross Sea, which are
connected by a U-shaped cyclonic feature in the vicinity of
the Ross Ice Shelf.
In the Ross Sea region, several distinct water masses can
be distinguished and are generally defined by the particular
relationships between temperature and salinity (Jacobs and
Comiso 1989, 1992; Jacobs and Giulivi 1989; Carmack
1992). Three water masses can be distinguished in the
deeper oceanic waters of the Ross Sea and five over the
continental shelf and slope (Gouretski 1999).
Antarctic Surface Water, Circumpolar Deep Water,
and Antarctic Bottom Water occupy the oceanic parts of
the region. Of these, Circumpolar Deep Water constitutes
the greatest volume of water in the Southern Ocean and is
characterized by high nutrient levels, generated from
phytoplankton as it sinks through the water column.
Antarctic Bottom Water occupies the deepest region of
the water column and plays a major role in the global
oceanic system.
Antarctic Surface Water, High Salinity Shelf Water, Low
Salinity Shelf Water, Ice Shelf Water, and modified Circum-
polar Deep Water can be identified over the continental shelf
and slope of the Ross Sea, although in contrast to other water
masses, those of the continental shelf are highly variable.
Antarctic Surface Water is found throughout the Ross Sea
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Biology of the Southern Ocean
during the summer, with surface temperatures generally
higher in the south where the pack ice opens up earlier in
the summer and a wide range of salinities occur. Below this
layer are dense shelf waters. High Salinity Shelf Water is the
densest water found in the Southern Ocean and these high
salinities are explained by the intensive processes of ice
formation in polynas along the coast (Jacobs, Fairbanks, and
Horibe 1985). Low Salinity Shelf Water is found in the
eastern Ross Sea and it is marked by slightly warmer sea
surface freezing temperatures, and low salinities. Ice Shelf
Water exhibits temperatures below the sea surface freezing
point, an indication of interaction with the Ross Ice Shelf at
depth. This water is concentrated in the west of the Ross Sea
where it emerges from under the ice shelf (Jacobs and Giuvili
1999). Modified Circumpolar Deep Water represents deep
oceanic waters that have penetrated the continental shelf.
This water is warmer and occupies depths between 100
and 300 m.
1.8 BOTTOM WATER FORMATION
Antarctic Bottom Water is an important component of the
Southern Ocean system, responsible for about 34 Sv of the
northward flow, and providing cold water at depth to all
the oceans of the southern hemisphere. Its formation involves
a mixing of water types, helped by surface interaction.
Observations from the Discovery Expeditions (Deacon
1937) confirmed the Weddell Sea as the main source of
Bottom water. It accounts for more than 50% of the total
volume produced. Second to this is the Ross Sea Shelf
(Jacobs, Amos, and Buckhausen 1970; Jacobs, Fairbanks,
and Horibe 1985) with some other sites in the sector from
308E to 1708E also being possible sources (Baines and
Condie 1998).
According to Gordon (1988), postulated processes
and sources contributing to Weddell Sea Bottom Water
formation include haline convection by evaporation or
freezing in open leads and polynas, cooling under the
Filchner Ice Shelf, Ekman-layer effects, sinking along
frontal zones, derivation form deeper oceanic areas,
overflow of dense water from the Bransfield Strait, and
soluble diffusive convection. Estimates of the circumpolar
production rates of Bottom Water are in excess of 13 Sv
(Jacobs, Fairbanks, and Horibe 1985). The principal
mechanism was believed to involve frontal zone mixing
between water masses at the continental shelf break, with
the dense water sinking down the slope (Gill 1973;
Foster and Carmack 1976). Recent observations by
Norwegian scientists (Foldvik and Gammelsrad 1988)
have shown that large volumes of Weddell Sea Bottom
Water originate from Ice Shelf Water formed under
floating ice shelves by flowing down the continental
slope as organized plumes and mixing with the overlying
Weddell Sea Deep Water.
 The Southern Ocean
1.9 NUTRIENTS
In the Southern Ocean the concentration of nutrients is
lowest in the surface waters and greatest in the Warm
Deep Water for nitrate and phosphate, and usually near the
bottom for silicate. In general, the concentration of nutrients
in surface waters south of the Polar Front is much higher than
that found in other oceanic waters (Knox 1970; El-Sayed
1978). Nutrient rich water upwells at the Antarctic Diver-
gence spreading out to ultimately downwell at the Polar
Front. On the basis of data from other upwelling regions,
with time one would expect to find almost complete stripping
of plant nutrients in the upper 50–100 m due to the growth of
phytoplankton (Barlow 1982). Holm-Hansen (1985) esti-
mated that between 25 and 30 mM of nitrate per liter would
have been consumed by the time the upwelled water reached
the zone of the Polar Front (approximately 200 days in
3394—CHAPTER 1—25/10/2006—14:52—KADAMBADI—14440—XML MODEL C – pp. 1–15
15
transit), and postulated that one would expect to find
almost complete stripping of the nutrients in aged waters
close to the Polar Front. However, data from studies by
Kuramoto and Koyama (1982), and Watanabe and Nakajima
(1982) from a transect along 458 longitude reveals a
completely different picture. Nitrate was 27 mM at 628S,
22 mM at the Polar Front (528S) and 20 mM a few degrees
south of the Subtropical Convergence (418S). Silica, on the
other hand, was high (57 mM) at 678S and it decreased
sharply to 15 mM at 608S, and dropped to low values
(approximately 1 mM) just north of the Polar Front, reflecting
its assimilation and polymerization by diatoms and silico-
flagellates. The significance of these data and the degree to
which nutrient depletion can occur in the waters of the
Southern Ocean will be considered in detail in Chapter 2
(Figure 1.10).
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