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MARINE
> /~\

Edited by Fiona J. Scott

& Harvey J. Marchant
 Digitized by the Internet Archive
in 2013

http://archive.org/details/antarcticmarinepOOfion
ANTARCTIC MARINE PROTISTS
ANTARCTIC MARINE
PROTISTS
Edited by Fiona J. Scott
& Harvey J. Marchant

AUSTRALIAN BIOLOGICAL RESOURCES STUDY, CANBERRA

AUSTRALIAN ANTARCTIC DIVISION, HOBART
 © Commonwealth of Australia 2005
This work is copyright. Apart from any use as permitted under the Copyright Act 1968, no part may be

reproduced or distributed by any process or stored in any retrieval system or data base without prior
written permission from the copyright holder. Requests and inquiries concerning reproduction and
rights should be addressed to the Director, Australian Biological Resources Study, GPO Box 787,
Canberra, ACT 2601, Australia, or to the Director, Australian Antarctic Division, Channel Highway,
Kingston, Tasmania 7050, Australia.

This book is available from:

Australian Biological Resources Study

GPO Box 787
Canberra ACT 2601
Australia

Fax: (02)6250 9555 Int: +(61 2) 6250 9555

email: abrsweb@deh.gov.au

Australian Biological Resources Study is a program within the Parks Australia Division of the
Australian Government's Department of the Environment and Heritage.

Australian Antarctic Division is part of the Australian Government's Department of the Environment
and Heritage.

National Library of Australia

Cataloguing-in-Publication entry

Antarctic Marine Protists

Bibliography.
Includes index.
ISBN 642 56835 9

1. Protists - Marine protozoa -

Identification. 2.
Antarctica - Microalgae - Antarctica -
Identification. 3.

Identification. I. Scott, Fiona Jean. II. Marchant, Harvey John.

III. Australian Biological Resources Study. IV. Australian Antarctic Division.

579

Cover design and digital illustration by Vanessa Tucker

Published by ABRS, Canberra and AAD, Hobart

Printed by Goanna Print, Canberra
 . . 7

CONTENTS

Contributors VI

Acknowledgements vii

1 Introduction, Harvey J. Marchant & Fiona J. Scott 1

Systematic Arrangement of Taxa 5

2. Diatoms, Fiona J. Scott & David P. Thomas 13

3. Dinoflagellates, Andrew McMinn & Fiona J. Scott 202

4. Silicoflagellates, Gustaaf M. Hallegraeff 251
5. Haptophytes: Order Prymnesiales,
Harvey J. Marchant & Fiona J. Scott & Andrew T. Davidson 255
6. Haptophytes: Order Coccolithophorales,
Claire S. Findlay, Jeremy R. Young & Fiona J. Scott 276
7. Chrysophytes, Harvey J. Marchant & Fiona J. Scott 295
8. Prasinophytes, Harvey J. Marchant 308
9. Chlorophytes, Fiona J. Scott 316
10. Cryptophytes, Fiona J. Scott & John van den Hoff 3 1

1 1 Euglenoids, Fiona J. Scott 319

12. Cyanophytes, Harvey J. Marchant 324
13. Choanoflagellates, Harvey J. Marchant 326
14. Ciliates, Wolfgang Petz 347
15. Protista Incertae Sedis, Fiona J. Scott & Harvey J. Marchant 449

Glossary 470
Bibliography 489
Copyright Permission 542
Index 549
 CONTRIBUTORS

Ms Fiona J. Scott, Australian Antarctic Division, Channel Highway, Kingston, Tasmania

7050, Australia

Dr Harvey J. Marchant, Australian Antarctic Division, Channel Highway, Kingston,

Tasmania 7050, Australia

Dr Andrew T. Davidson, Australian Antarctic Division, Channel Highway, Kingston,

Tasmania 7050, Australia
Dr Claire S. Findlay, c/o Australian Antarctic Division, Channel Highway, Kingston,
Tasmania 7050, Australia
Assoc. Prof. Gustaaf M. Hallegraeff, Department of Plant Sciences, University of
Tasmania, Churchill Avenue, Sandy Bay, Tasmania 7005, Australia

Dr Andrew McMinn, IASOS, University of Tasmania, Churchill Avenue, Sandy Bay,

Tasmania 7005, Australia
Dr Wolfgang Petz, Institute of Zoology, University of Salzburg, Hellbrunnerstrasse 34, A-
5020, Salzburg, Austria, and TB Petz, Ecological Consulting Office, Hallwanger Landes-
strasse 32a, A-5300 Hallwang, Austria
Prof. Patrick G. Quilty, Department of Earth Sciences, University of Tasmania, Churchill
Avenue, Sandy Bay, Tasmania 7005, Australia

Dr David P. Thomas, Department of Plant Sciences, University of Tasmania, Churchill

Avenue, Sandy Bay, Tasmania 7005, Australia

Dr Paul G. Thomson, Australian Antarctic Division, Channel Highway, Kingston, Tasmania

7050, Australia

Mr John van den Hoff, Australian Antarctic Division, Channel Highway, Kingston,
Tasmania 7050, Australia

Dr Jeremy R. Young, Department of Palaeontology, The Natural History Museum,

Cromwell Road, London SW7 5BD, United Kingdom

VI
 ACKNOWLEDGEMENTS
Much of the preparation of this volume was supported by funding initially from a Marine
Sciences and Technologies grant and, subsequently, from the Australian Biological Resources
Study. The Australian Antarctic Division of the Australian Government's Department of the
Environment and Heritage provided logistical support for the collection of material, as well as
laboratory, office and staff support during the final two years of research.
Many individuals, including Leanne Armand, Sue Blackburn, Pru Bonham, Susan Campbell,
Vicki Cochrane, Roger Croome, Laura Cunningham, Allan Davey, Stella Humphries, Akira
Ishikawa, Jiang Jialun, Richard Norris, Lu Peiding, Rick Perrin, Sandra Potter, Pat Quilty,
John Shearer, Michael Stoddart, Ron Thompson, Paul Thomson, Naja Vors, Tim Walker and
Simon Wright, have contributed in various ways to its production; our sincere thanks go to
them. Special thanks go to Ms Geraldine (Gerry) Nash of the Electron Microscope Unit,
Australian Antarctic Division. Her support and encouragement throughout the project has
been instrumental in the production of this volume. We are most grateful to Vanessa Tucker
for her creative digital illustration and design of the cover.

Photographs and images are the works of the chapter authors and Australian Antarctic
Division staff (John Cox, Andrew McEldowney, Anna McEldowney, Sandy Molloy, Gerry
Nash, Rick Perrin, Rick van den Enden, Esmee van Wijk, Karen Westwood) unless
specifically credited in the captions or the Copyright Permission section of the book.
A number of previously published images are also included. These are acknowledged in the
Copyright Permission section near the end of the book.

The authors of Chapter 6 thank Dr Joerg Bollmann (Swiss Federal Institute of Technology,
Zurich) for permission to reproduce images and for his assistance in accessing the EMIDAS
database (Bollman et al., 1999), and Michael Knappertsbusch and Sabrina Renaud for
permission to use their images.

PatrickMcCarthy was the ABRS project manager and editor. Several current and former
ABRS members also assisted: Virginnia Du Toit, Katy Mallett, Brigitte Kuchlmayr,
staff
Helen Thompson and Tony Orchard.

vn
 1. INTRODUCTION
Harvey J. Marchant & Fiona J. Scott

This volume is the result of a long-recognised need for a guide to the pelagic eukaryotic
microorganisms of the Antarctic marine environment for use principally by non-specialists.
It was originally conceived as an in-house handbook for our studies on phytoplankton, but in

the absence of a synthesis of the taxonomy of Antarctic marine protists and with the
encouragement of other Antarctic researchers with similar needs, we have expanded the
original concept.

Our aim has been provide an illustrated description for each formally described species of
to
Antarctic marine protist. In addition, we have attempted to collate the widely scattered
literature on the taxonomy of Antarctic pelagic protists in order to resolve some of the
taxonomic confusion that has arisen, especially with regards to synonymy.

The Kingdom Protista comprises those eukaryotic organisms historically and collectively
known as phytoplankton as well as the diverse groups of non-photosynthetic organisms, the
protozoa, although many zoological texts include photosynthetic eukaryotes as protozoa.
Here we include the pelagic eukaryotic protists, in addition to the cyanobacterium
Synechococcus, which are found at and south of the Polar Front.

The Polar Front, formerly known Convergence, represents the major circum-
as the Antarctic
polar oceanographic boundary between warmer water to the north and colder
sinking
southern water. Water temperature differs by about 2°C across the Front, one of the most
significant oceanographic boundaries on Earth. The region south of the Polar Front can be
divided into three zones: the Permanently Open Ocean Zone; the Seasonal Ice Zone; and the
Coastal and Continental Shelf Zone (Fig. 1.1). Because of the major environmental differ-
ences between these zones, their biology differs dramatically.

In addition to the water column, sea-ice is an important habitat for marine protists. From
about March each year, the sea surface around Antarctica freezes, reaching a maximum
extent in September when nearly 20 million square kilometres are ice-covered to an average
thickness of slightly less than a metre. Sea-ice has a highly complex structure presenting a
great diversity of habitats for microorganisms. For further information on the structure of the
Southern Ocean and sea-ice as a habitat for protists, see El-Sayed & Fryxell (1993), Garrison
& Gowing (1993), Marchant & Murphy (1993), Palmisano & Garrison (1993), Knox (1994),
Fogg (1998) and Thomas & Dieckmann (2003).
Antarctic marine protists are amazingly diverse. They span a size range from around 2 p.m in
some Parmales to several millimetres long in some diatoms. Their shapes are highly variable,
and many bear flagella or cilia to enable movement or the generation of water currents to
assist in feeding. Some are encased in rigid walls of silica, cellulose or other structural
components, while others are covered in scales or mucus. The life cycles of some protists are
comparatively simple, while in others they are so complex as to lead to taxonomic confusion.

Autotrophic protists employ a wide array of pigments in addition to chlorophyll a. This

diversity and that of lipids and other chemical constituents have been exploited to
characterise and quantify the abundance of the various microalgal groups (Jeffrey et al.,
1997). The range of feeding strategies of heterotrophic protists is also considerable, and this
can lead to a blurring of the distinction between plants and animals. Several autotrophs can
feed heterotrophically, and several heterotrophs, most commonly ciliates, can retain and
nurture the chloroplasts of ingested prey to supplement their nutrition. This process
(mixotrophy) is not confined to protists; it is also employed by sponges, corals and even
some higher plants. Detailed discussion of the biology of protists is beyond the scope of this
work, and readers are directed to the recent review by Hausmann et al. (2003).

Antarctic marine phytoplankton, particularly diatoms, have received considerable attention

since the earliest explorations of the Antarctic. Hooker's collections from the Erebus and
Terror expeditions of 1839-43, examined by Ehrenberg, formed the basis of the first paper
 INTRODUCTION

on Antarctic phytoplankton (Ehrenberg, 1844b). The development of our knowledge of

Antarctic marine protists has been traced by several authors (e.g. Vincent, 1988; Medlin &
Priddle, 1990; Karl, 1993; Knox, 1994) and, specifically in the Australian Antarctic program,
by Marchant et al. (2002).
own observations, we have drawn on other published sources in attempting
In addition to our
to incorporate as many of the reported Antarctic marine protists as possible. Apart from
those organisms that have been formally described, for completeness we have also included
some undescribed taxa (as type A, B, C, etc.). We do not describe any new taxa.
For phytoplankton species, author abbreviations follow R.K.Brummitt & C.E.Powell,
Authors of Plant Names (Royal Botanic Gardens, Kew, 1992). Journal titles are abbreviated
in accordance with G. H.M.Lawrence et al., Botanico-Periodicwn-Huntianum (Hunt
Botanical Library, Pittsburgh, 1968) and G.D.R.Bridson & E.R.Smith, Botanico-Periodicum-
Huntianum/Supplementum (Hunt Institute for Botanical Documentation, Pittsburgh, 1991),
or, if not included in those works, are abbreviated in the same style. Other literature is
abbreviated in accordance with F.A.Stafleu & R.S. Cowan, Taxonomic Literature, 2nd edn
(Bohn, Scheltema & Holkema, Utrecht, 1976-87), except that upper case initial letters are
used for proper names and significant words.

While we have attempted to make this treatment as comprehensive as possible, we are well
aware that our coverage of organisms is uneven. Thus, the accounts of, for example, diatoms
and choanoflagellates are more comprehensive than those of prasinophytes and amoebae.
This is a consequence of the uneven attention given to the various groups, either due to their
overwhelming abundance and/or because of their localised importance within communities.
Similarly, while we have tried to refer to all of the relevant literature, it is likely, despite our
best efforts, that we have missed material especially in the 'grey' literature and publications
in languages other than English. We trust that this book will be seen as an attempt to
assemble descriptive information on the Antarctic marine protists, and thus provide an
overview of current knowledge. We hope that the information provided here will stimulate
the discovery and documentation of new records and taxa.

Perhaps most importantly, with the rapidly increasing use of molecular techniques to identify
protists, thisvolume will provide a comprehensive treatment of protist morphology which
can be used in conjunction with molecular studies.

Why are Antarctic marine protists important?

Before the 1980s, the Southern Ocean around Antarctica was thought to be one of the richest
marine ecosystems on Earth with one of the simplest food webs in which diatoms were
grazed by krill, and krill were consumed by fish, seals, birds and whales. We now know that
this is a gross oversimplification and that while the Southern Ocean does have some highly
productive areas, in overall terms it is no more productive than the nutrient-poor parts of the
world's oceans, with food webs that are no less complex than those of lower latitudes. The
microbial components of Antarctic food webs form marine snow —
amorphous macroscopic
aggregates of microorganisms in a matrix of organic and inorganic detritus —
and function
as a microbial loop similar to microorganisms elsewhere in the world's oceans. However,
there is one important difference, viz. the concentration of picoplanktonic cyanobacteria is
very low in Antarctic waters, and this has ramifications for the dynamics of the microbial
loop.

As well as forming the base of the food web on which all other marine organisms depend,
marine protists play a role in influencing climate. The Southern Ocean is a key area of
exchange of carbon dioxide with the atmosphere. The seasonal and spatial variation in the
microbial community composition is a principal determinant of whether the ocean is drawing
down atmospheric carbon dioxide or is a source of carbon dioxide to the atmosphere. These
processes, the so-called "biological pump", involve the conversion of inorganic carbon to
organic carbon by photosynthesis and the transport of the latter to the deep sea and sea-floor
(Moore & Bolin, 1987; Knox, 1994). Some of the highest rates of vertical flux of carbon in
the ocean have been measured in Antarctic waters. Highest drawdown of atmospheric carbon
 INTRODUCTION

South Georgia

120°W\
120°E

150°W
150°E

Permanent Ice Shelves

Permanent Open Ocean Zone

Seasonal Ice Zone

Coastal and Continental Shelf Zone

Figure 1.1. Biological zones in the Southern Ocean, with the main collecting sites for marine
protists.
 INTRODUCTION

dioxide is usually observed during spring or early summer at the times of dramatic blooms of
phytoplankton, especially diatoms. At other times of the year microbial respiration supplies
carbon dioxide to the atmosphere as well as having a pivotal role in carbon dioxide
biogeochemistry.

Antarctic protists produce and modify the concentrations of reduced sulphur compounds,
especially dimethyl sulphide (DMS). When DMS
is ventilated to the atmosphere it produces

aerosol particles that promote the formation of clouds. Clouds play a significant role in
global climate by reflecting the Sun's energy back into space. The Southern Ocean is
reportedly the source of approximately 17 per cent of the total global production of these
sulphur compounds (Curran &
Jones, 2000). The role of the Antarctic protistan community
composition in global biogeochemical processes and their climatic consequences cannot be
overstated.

Sample collection and processing

Because marine protists include such a diverse range of organisms, no single technique can
be employed to sample them. For the purposes of this volume, organisms were collected in a
variety of ways from the different depths of the water column. Surface seawater samples
were collected either by a bucket or by centrifugal pump, and subsurface samples were
collected using various types of bottles. Organisms were concentrated onto polycarbonate
filters (pore size 0.2-8.0 um) by gentle filtration or by continuous flow centrifugation. Some
specimens, including the larger diatoms, were collected by net hauls with 20 um mesh nets,
or by pumping surface water through 20 um mesh netting.

In many cases enrichment cultures of organisms were established at sea from which clonal
cultures of specific organisms were later established. Both enrichment and clonal cultures
were maintained in GP5 of G medium; Loeblich & Smith, 1968), f/2
(a modification
(Guillard & Ryther, 1962) or MET
(Schone & Schone, 1982) media. Concentrated living
material was identified, enumerated, photographed or video-taped using a light microscope
and associated equipment isolated from the vibrations of the ship (Marchant, 1985a). The
shipboard laboratories were maintained at temperatures of less than 5°C, or the microscope's
stage was cooled to enable examination of the morphology and/or swimming patterns of the
motile species and those species that do not preserve well in standard fixatives.

On litre surface water samples were fixed and stored in 1% glutaraldehyde

various cruises, 2
in preparation for scanning and/or transmission electron microscopy (Johnson & Sieburth,
1982). This technique is useful for preserving delicate cells including those infected with
viruses. Concentrated cells were also fixed for subsequent light and electron microscopy.
The most commonly used technique was the fixation of organisms in 1% glutaraldehyde in
buffered seawater, maintaining this material at about 4°C until further examination.

For transmission electron microscopy (TEM) of scale-covered flagellates and whole mounts
of choanoflagellates, concentrated cells were fixed in osmium tetroxide vapour, either dried
onto Formvar-coated grids or allowed to settle onto polylysine-treated Formvar-coated grids
(Marchant & Thomas, 1983). Grids were then washed with distilled water, dried and
subsequently negative-stained with uranyl acetate or shadow cast with platinum/palladium or
chromium for TEM examination. For TEM of sectioned material, glutaraldehyde-fixed
organisms were post-fixed with osmium tetroxide, dehydrated through a graded acetone
series before being embedded in Spurr's (1969) low viscosity resin. Thin sections were cut
with either glass or diamond knives, and the sections were stained with uranyl acetate and
lead citrate.

Scanning electron microscopy (SEM) was performed on glutaraldehyde-fixed cells that had
settled on polylysine-coated microscope coverglasses, dehydrated with a graded series of
acetone, critical point-dried and sputter-coated with gold. In order to preserve the scales of
coccolithophorids, 0.8 um pore-size polycarbonate filters, onto which cells had been
concentrated, were rinsed with freshly prepared distilled water and rapidly air-dried. They
were sputter-coated with gold and examined by SEM without any other treatment.
 INTRODUCTION

Protargol, the most common silver-impregnation technique for ciliates, is generally well-
suited for marine species. In the modification of Wilbert (1975), specimens are fixed in
Bouin or Stieve's fluid (mercuric chloride sublimate) in staining blocks, washed in distilled
water, bleached with dilute aqueous sodium hypochlorite, washed again, impregnated in 1%
protargol (silver protein) solution, developed in diluted 1% hydroquinone solution, fixed in
2.5% sodium thiosulphate, washed again in distilled water, mounted with albumen-glycerol
(1:1 egg albumen and glycerol) onto a microscope slide, dried, dehydrated with a graded
series of alcohol, cleared twice in xylene and mounted in a synthetic, neutral mounting
medium (e.g. Permount, Eukitt or Entellan). Optimal dilutions and treatment times depend on
the species, and these have to be determined empirically in advance. Several other protocols
are also available (e.g. Montagnes & Lynn, 1987; Foissner, 1991; Lee & Soldo, 1992).

SYSTEMATIC ARRANGEMENT OF TAXA

CLASS DIATOMOPHYCEAE/ Coscinodiscus centralis
BACILLARIOPHYCEAE Coscinodiscus curvatulus
Order Centrales Coscinodiscus horridus
Coscinodiscus marginatus
Family Asterolampraceae
Coscinodiscus oculoides
Aster omphalus heptactis
Coscinodiscus oculus-iridis
Asteromphalus hookeri
Coscinodiscus oppositus
Asteromphalus hyalinus
Coscinodiscus radiatus
Asteromphalus parvulus
Coscinodiscus subtilis
Asteromphalus roperianus
Stellarima microtrias
Eucampia antarctica var. antarctica
Eucampia antarctica var. recta Family Eupodiscaceae
Trigonium arcticum Odontella aurita
Odontella litigiosa
Family Chaetoceraceae
Odontella weissflogii
Chaetoceros adelianum
Chaetoceros aequatorialis Family Heliopeltaceae
var. antarctica Actinoptychus senarius
Chaetoceros atlanticus Family Hemidiscaceae
Chaetoceros brevis Actinocyclus actinochilus
Chaetoceros bulbosus Actinocyclus adeliae
Chaetoceros castracanei Actinocyclus curvatulus
Chaetoceros concavicornis Actinocyclus divisus
Chaetoceros convolutus Actinocyclus exiguus
Chaetoceros criophilus Actinocyclus octonarius
Chaetoceros curvatus Actinocyclus spiritus
Chaetoceros curvisetus Azpeitia nodulifera
Chaetoceros debilis Azpeitia tabularis
Chaetoceros dichaeta Hemidiscus cuneiformis
Chaetoceros flexuosus Roperia tesselata
Chaetoceros gaussii Family Leptocylindraceae
Chaetoceros hendeyi Corethron inerme
Chaetoceros laciniosus Corethron pennatum
Chaetoceros neglectus Leptocylindrus danicus
Chaetoceros neogracilis Leptocylindrus mediterraneus
Chaetoceros pendulus
Family Melosiraceae
Chaetoceros peruvianus
Melosira adeliae
Chaetoceros radicans
Melosira sol
Chaetoceros simplex
Family Rhizosoleniaceae
Chaetoceros socialis
Dactyliosolen antarcticus
Chaetoceros tortissimus
Dactyliosolen tenuijunctus
Family Coscinodiscaceae Guinardia cylindrus
Coscinodiscus asteromphalus Guinardia tubiformis
Coscinodiscus bouvet Proboscia alata
 INTRODUCTION

Proboscia inermis Amphora type A

Proboscia truncata Amphora type B
Rhizosolenia antennata f. semispina Amphora type C
Rhizosolenia chunii Family Entomoneidaceae
Rhizosolenia crassa Amphiprora kufferathii
Rhizosolenia curvata Entomoneis kjellmanii
Rhizosolenia hebetata f. semispina Entomoneis paludosa var. hyperborea
Rhizosolenia imbricata
Family Fragilariaceae
Rhizosolenia polydactyla
Fragilaria islandica var. islandica
f. polydactyla
Fragilaria islandica var. adeliae
Rhizosolenia rhombus
Fragilaria islandica var. producta
Rhizosolenia sima f. sima
Fragilaria islandica var. stricta
Rhizosolenia simplex
Grammatophora arcuata
Rhizosolenia sty I [form is
Pseudostaurosira brevistriata
Family Thalassiosiraceae Synedropsis fragilis
Lauderia annulata Synedropsis hyperboreoides
Planktoniella sol Synedropsis laevis
Porosira glacialis Synedropsis lata
Porosira pseudodenticulata Synedropsis recta
Thalassiosira ambigua Thalassionema nitzschioides
Thalassiosira antarctica Thalassiothrix antarctica
Thalassiosira australis Trichotoxon reinboldii
Thalassiosira dichotomica
Family Naviculaceae
Thalassiosira eccentrica
Banquisia belgicae
Thalassiosira ferelineata
Berkeleya rutilans
Thalassiosira frenguellii
Ephemera planamembranacea
Thalassiosira frenguelliopsis
Gyros igma sp.
Thalassiosira gerloffii
Haslea trompii
Thalassiosira gracilis var. gracilis
Luticola muticopsis
Thalassiosira gracilis var. expecta
Manguinea fusiformis
Thalassiosira gravida
Membraneis challengeri
Thalassiosira lentiginosa
Navicula criophila
Thalassiosira leptopus
Navicula cristata
Thalassiosira maculata
Navicula directa var. directa
Thalassiosira margaritae
Navicula gelida var. parvula
Thalassiosira oestrupii
Navicula glaciei
Thalassiosira oliveriana
Navicula gracilis var. antarctica
Thalassiosira partheneia
Navicula schefterae
Thalassiosira perpusilla
Pinnularia quadratarea
Thalassiosira poro-irregulata
var. constricta
Thalassiosira poroseriata
Plagiotropus gaussii
Thalassiosira ritscheri
Pleurosigma directum
Thalassiosira scotia
Pseudogomphonema kamtschaticum
Thalassiosira symmetrica
Trachyneis aspera
Thalassiosira trifulta
Family Auriculaceae
Thalassiosira tumida
Auricula compact a
Order Pennales
Family Bacillariaceae
Family Achnanthaceae
Cylindrotheca closterium
Achnanthes brevipes
Diploneis stigmosa
Achnanthes vicentii
Fragilariopsis curta
Cocconeis costata var. antarctica
Fragilariopsis cylindrus
Cocconeis fasciolata
Fragilariopsis kerguelensis
Cocconeis infirmata
Fragilariopsis linearis
Cocconeis schitettii
Fragilariopsis obliquecostata
Cocconeis type A
Fragilariopsis pseudonana
Family Cymbellaceae Fragilariopsis rhombica
Amphora racovitzae Fragilariopsis ritscheri
 INTRODUCTION

Fragilariopsis separanda Order Suessiales

Fragilariopsis sublinearis Family Suessiaceae
Fragilariopsis vanheurckii Polarella glacialis
Nitzschia acicularis
Order Gonyaulacales
Nitzschia australis
Family Ceratiaceae
Nitzschia barbieri
Ceratium extensum/C. fusus complex
Nitzschia bicapitata
Ceratium pentagonum/C. lineatum
Nitzschia decipiens
complex
Nitzschia lecointei
Ceratium tripos
Nitzschia longissima
Nitzschia medioconstricta Family Goniodomataceae
Nitzschia peragallii Alexandrium tamarense
Nitzschia sicula var. bicuneata Family Gonyaulacaceae
Nitzschia stellata Gonyaulax digitale
Nitzschia taenia Gonyaulax scrippsae
Pseudonitzschia antarctica Gonyaulax striata
Pseudonitzschia heimii Family Oxytoxaceae
Pseudonitzschia lineola Oxytoxum criophilum
Pseudonitzschia prolongatoides
Order Peridinales
Pseudonitzschia subcurvata
Family Calciodinellaceae
Pseudonitzschia turgidula
Scrippsiella troichoidea
Pseudonitzschia turgiduloides
Family Kolkwitziellaceae
CLASS DINOPHYCEAE Diplopsalis lenticula
Preperidinium meunierii
Order Prorocentrales
Preperidinium perlatum
Family Prorocentraceae
Family Podolampaceae
Mesoporous perforatus
Blepharocysta splendor-maris
Prorocentrum antarcticum
Podolampas antarctica
Prorocentrum balticum
Prorocentrum compressum Family Protoperidiniaceae
Protoperidinium adeliense
Order Dinophysales
Protoperidinium affine
Family Amphisoleniaceae Protoperidinium antarcticum
Dinophysis antarctica
Protoperidinium applanatum
Dinophysis contracta Protoperidinium archiovatum
Dinophysis cornuta
Protoperidinium areolatum
Dinophysis meteori
Protoperidinium bellulum
Dinophysis operculoides Protoperidinium bipatens
Dinophysis ovum
Protoperidinium charcoti
Dinophysis punctata
Protoperidinium concavum
Dinophysis tenuivelata Protoperidinium crassipes
Dinophysis tuberculata
Protoperidinium cruciferum
Heteroschisma subantarctica
Protoperidinium curium
Phalacroma rotundatum Protoperidinium defectum
Order Gymnodiniales Protoperidinium elegantissimum
Family Gymnodiniaceae Protoperidinium glyptopterum
Amphidinium hadai Protoperidinium granii
Gymnodinium baccatum Protoperidinium incertum
Gymnodinium frigidum Protoperidinium incognitum
Gymnodinium modestum Protoperidinium latistriatum
Gymnodinium soyai Protoperidinium mediocre
Gyrodinium glaciale Protoperidinium melo
Gyrodinium lachryma Protoperidinium metananum
Gyrodinium rhabdomante Protoperidinium penitum
Katodinium sp. Protoperidinium rosaceum
Torodinium sp. Protoperidinium thulesense
Family Polykrikaceae Protoperidinium unipes
Polykrikos schwartzii
 INTRODUCTION

CLASS DICTYOCHOPHYCEAE Family Syracosphaeraceae

Order Dictyochales Calciopappus sp. (HET)
Syracosphaera molischii (HET)
Family Dictyochaceae
Syracosphaera pulchra (HET, HOL)
Dictyocha fibula
Dictyocha octonaria Family Umbellosphaeraceae
Dictyocha speculum Umbellosphaera irregularis (HET)
Genera Incertae Sedis
CLASS HAPTOPHYCEAE/ Trigonaspis melvillea (HOL)
PRYMNESIOPHYCEAE Wigwamma annulifera (HET, HOL)
Order Prymnesiales Wigwamma antarctica (HET)

Family Phaeocystaceae
Wigwamma arctica (HET)
Phaeocystis antarctica
Wigwamma triradiata (HET)

Family Prymnesiaceae
CLASS CHRYSOPHYCEAE
Chrysochromulina acantha
Order Ochromonadales
Chrysochromulina bergenensis
Chrysochromulina chiton Family Paraphysomonadaceae
Chiysochromulina ericina Paraphysomonas antarctica
Chrysochromulina hirta Paraphysomonas butcheri
Chiysochromulina leadbeateri Paraphysomonas cribosa
Chrysochromulina polylepis Paraphysomonas cylicophora
Chrysochromulina vexillifera Paraphysomonas imperforata
Chrysochromulina type A Paraphysomonas oligocycla
Chiysochromulina type B Paraphysomonas vestita

Chrysochromulina type C Paraphysomonas type A

Chrysochromulina type D Order Parmales
Chrysochromulina type E Family Triparmaceae
Chrysochromulina type F Tetraparma pelagica
Chrysochromulina type G Triparma columacea subsp. a lata
Chiysochromulina type H Triparma laevis subsp. pinnatilobata
Chrysochromulina type I Triparma laevis subsp. ramispina
Chrysochromulina type J Triparma strigata
Chrysochromulina type K
Family Pentalaminaceae
Chrysochromulina type L
Pentalamina corona
Chrysochromulina type M Order Synurales
Order Coccolithophorales
Family Pleurochloridaceae
Family Calcidiscaceae
Meringosphaera mediterranea
Calcidiscus leptoporus (HET, HOL)*
Family Helicosphaeraceae CLASS PRASINOPHYCEAE
Helicosphaera carteri var. carteri
Order Chlorodendrales
(HET, HOL)
Family Halosphaeraceae
Family Noelaerhabdaceae Pterosperma parallelum
Emiliania huxleyi (HET)
Pterosperma cf. polygonum
Gephyrocapsa caribbeanica (HET) Pyramimonas australis
Gephyrocapsa ericsonii (HET)
Pyramimonas gelidicola
Gephyrocapsa muellerae (HET) Pyramimonas tychotreta
Gephyrocapsa oceanica (HET)
Order Mamiellales
Family Papposphaeraceae
Pappomonas weddellensis (HET) Family Mamiellaceae
Papposphaera arctica (HET, HOL) Mantoniella antarctica
Papposphaera borealis (HET, HOL) Mantoniella squamata
Papposphaera obpyramidalis (HET)
Papposphaera polybotrys (HET, HOL) CLASS CHLOROPHYCEAE
Papposphaera simplicissima (HET) Order Volvocales
Family Chlamydomonaceae
Polytoma papillata
HET, heterococcolithophore; HOL,
holococcol ithophore
 INTRODUCTION

CLASS CRYPTOPHYCEAE Stephanoeca diplocostata

Order Cryptomonadales var. paucicostata
Stephanoeca norrisii
Family Cryptomonadaceae
Geminigera cryophila
cryptomonad indet.
PHYLUM CILIOPHORA
CLASS HETEROTRICHEA
CLASS EUGLENOPHYCEAE Order Heterotrichida
Order Sphenomonadales Family Folliculinidae
Ascobius simplex
Family Sphenomonaceae
Folliculina ampulla
Anisonema prosgeobium
Lagotia expansa
Order Eutreptiales
Lagotia faurefrem ieti
Family Eutreptiaceae Lagotia gigantea
Eutreptiella cf. braarudii Lagotia similis
Eutreptiella sp. Parafolliculina amphora
Postgaardi mariagerensis Parafolliculina patagonica
Parafolliculina violacea
CLASS CYANOPHYCEAE Pseudofolliculina melitta
Order Chroococcales
Family Stentoridae
Synechococcus sp.
Heterostentor coeruleus
Family Condylostomatidae
CLASS CRASPEDOPHYCEAE/
Condylostoma granulosum
CHOANOFLAGELLIDA
Condylostoma remanei
Order Choanoflagellida
Family Codonosigidae CLASS SPIROTRICHEA
Monosiga ovata
Order Euplotida
Family Acanthoecidae
Family Uronychiidae
Acanthocorbis tintinnabulum
Diophrys appendiculata
Acanthocorbis unguiculata
Diophrys oligothrix
Acanthoeca brevipoda
Diophrys scutum
Acanthoeca spectabilis
Uronychia transfuga
Acanthoecopsis asymmetrica
Family Aspidiscidae
Bicosta antennigera
Aspidisca antarctica
Bicosta spinifera
Aspidisca crenata
Calliacantha natans
Aspidisca polypoda
Calliacantha simplex
Aspidisca quadrilineata
Cosmoeca norvegica
Aspidisca terranovae
Cosmoeca takahashii
Cosmoeca ventricosa Family Gastrocirrhidae
Crinolina aperta Cytharoides balechi
Diaphanoeca grandis Family Euplotidae
Diaphanoeca midtiannulata Euplotes acanthodus
Diaphanoeca pedicellata Euplotes antarcticus
Diaphanoeca sphaerica Euplotes balteatus
Kakoeca antarctica Euplotes euryhalinus
Parvicorbicula circularis Euplotes focardii
Parvicorbicula ongulensis Euplotes rariseta
Parvicorbicula socialis Euplotes sigmolateralis
Pleurasiga minima Order Tintinnida
Pleurasiga orculaeformis
Family Codonellidae
Pleurasiga reynoldsii
Tintinnopsis bacillaria
Polyfibula sphyrelata
Tintinnopsis buetschlii
Saepicula leadbeateri
Tintinnopsis lobiancoi
Spiraloecion didymocostatum
Tintinnopsis parva
Stephanoeca complexa
Tintinnopsis petasa
Stephanoeca cupula
Family Codonellopsidae
Codonellopsis balechi
 INTRODUCTION

Codonellopsis gaussi Rimostrombidium glacicolum

Codonellopsis glacialis Family Leegaardiellidae
Codonellopsis morchella Leegaardiella elbraechteri
Codonellopsis pusilla
Family Lohmanniellidae
Codonellopsis soyai
Lohmanniella oviformis
Laackmanniella naviculaefera
Order Urostylida
Stenosemella avellana
Stenosemella oliva Family Urostylidae
Stenosemella perpusilla Holosticha foissneri
Holosticha pullaster
Family Metacylididae
Holosticha spindleri
Cox lie I la frigida
Metaurostyla thompsoni
Coxliella intermedia
Metaurostylopsis rubra
Family Ascampbelliellidae Notocephalus parvulus
Acanthostomella norvegica
Family Pseudokeronopsidae
Family Epiplocylididae Thigmokeronopsis antarctica
Epiplocylis undella
Thigmokeronopsis crystallis
Epiplocyloides antarctica
Order Strombidiida
Family Ptychocylididae
Family Strombidiidae
Ptychocylis ostenfeldi
Laboea strobila
Ptychocylis wailesi
Spirostrombidium pseudocinctum
Family Rhabdonellidae Spirostrombidium rhyticollare
Protorhabdonella curta Strombidium acutum
Protorhabdonella simplex Strombidium antarcticum
Rhabdonella amor Strombidium conicoides
Family Petalotrichidae Strombidium crassulum
Parapetalotricha meridiana Strombidium diversum
Family Cyttarocylididae Strombidium elongatum
Cyttarocylis cassis Strombidium emergens
Family Xystonellidae Strombidium glaciale
Cymatocylis calyciformis Strombidium kryale
Cymatocylis convallaria Strombidium sulcatum
Cymatocylis cristallina Strombidium syowaensis
Cymatocylis drygalskii Strombidium tintinnodes
Cymatocylis kerguelensis Strombidium wulffi
Cymatocylis nobilis Tontonia antarctica
Cymatocylis vanhoeffeni
Protocymatocylis pseudoconica CLASS LITOSTOMATEA
Xy st on el I a acus Order Cyclotrichida
Family Undellidae Family Mesodiniidae
Undella claparedii Mesodinium pulex
Undella hyalina Myrionecta rubra
Family Tintinnidae Order Haptorida
Amphorellopsis quinquealata Family Didiniidae
Amphorides laackmanni Cyclotrichium gigas
Bursaopsis conicoides Didinium gargantua
Bursaopsis ollula Monodinium balbianii
Dadayiella ganymedes Family Pseudotrachelocercidae
Daturella frigida Pseudotrachelocerca trepida
Salpingella acuminatoides
Family Lacrymariidae
Salpingella costata
Lacrymaria lagenula
Salpingella decurtata
Lacrymaria spiralis
Order Choreotrichida
Family Acropisthiidae
Family Strobilidiidae Fuscheria marina
Pelagostrobilidium neptuni
Pelagostrobilidium spirale
Rimostrombidium conicum

10
 INTRODUCTION

Family Trachelophyllidae Order Philasterida

Chaenea teres Family Pseudocohnilembidae
Order Pleurostomatida Pseudocohnilembus pus ill us
Family Litonotidae Family Uronematidae
Kentrophyllum antarcticum Urocyclon ovatum
Litonotus antarcticus Uronema acutum
Litonotus emmerichi Uronema antarcticum
Litonotus kopimorphus Uronema elegans
Litonotuslamella Uronema marinum
Loxophyllum rostra turn Uronema paramarinum
Uropedalium antarcticum
CLASS PHYLLOPHARYNGEA Family Cryptochilidae
Order Chlamydodontida Cryptochilum reniforme
Family Kryoprorodontidae Family Philasteridae
Gymnozoum glaciate Porpostoma grassei
Gymnozoum sympagicum Porpostoma notatum
Gymnozoum viviparum Family Orchitophryidae
Family Lynchellidae Metanophrys antarctica
Chlamydonella prostomata Order Pleuronematida
Chlamydonella pseudochilodon Family Pleuronematidae
Order Dysteriida Pleuronema coronation
Family Dysteriidae Pleuronema glaciate
Aegyriana parol iva Pleuronema puytoraci
Dysteria calkins i Family Cyclidiidae
Dysteria monostyla Cyclidium varibonneti
Order Exogenida Paractedoctema acruosa
Family Ophryodendridae Order Sessilida
Ophryodendron conicum Family Vaginicolidae
Family Thecacinetidae Cothurnia antarctica
Thecacineta calix Cothurnia subglobosa
Order Endogenida
Family Acinetidae
PROTISTA INCERTAE SEDIS
Acineta grahami
PHYLUM AMOEBOZOA
Conchacineta rotunda
CLASS LOBOSEA
Pelagacineta campanula Order Amoebae
Pelagacineta interrupta Amoeba spp.

CLASS NASSOPHOREA CLASS FILOSEA

Order Synhymeniida Order Euglyphida

Family Orthodonellidae Family Paulinellidae

Orthodonella shenae Paulinella intermedia
Zosterodasys kryophilus
GROUP GRANULORETICULOSA
CLASS PROSTOMATEA CLASS FORAMINIFEREA
Order Prorodontida Order Foraminiferida
Family Placidae Family Globorotaliidae
Placus antarcticus Neogloboquadrina pachyderma
Family Colepidae
Tiarina antarctica
GROUP ACTINOPODA
CLASS PHAEODAREA
CLASS OLIGOHYMENOPHOREA CLASS HELIOZEA
Order Ciliophryida
Order Peniculida
Family Ciliophrydae
Family Frontoniidae
Actinomonas mirabilis
Fronton ia frigida
Pteridomonas danica

11
 INTRODUCTION

Order Centrohelida (Centroheliozoa) Order Bicosoecales

Acanthocystis paliformis Family Cafeteriaceae
Acanthocystis perpusilla Cafeteria roenbergensis
Acanthocystis turfacea
Heterophrys myriapoda SUBPHYLUM HETEROKONTA
Order Rotosphaerida Incertae Sedis
Pinaciophora denticulata Ancyromonas sigmoides
Pinaciophora fluviatilis
GENERA INCERTAE SEDIS
SUPERCLASS KINETOPLASTA Amastigomonas debruynei
CLASS BODONEA Cryothecomonas armigera
Order Kinetoplastida Metromonas simplex
Family Bodonida Petasaria heterolepis
Bodo curvifilus Telonema antarctica
Leucocryptos marina Telonema subtile
Thaumatomastix fragilis
PHYLUM CHROMISTA Thaumatomastix fusiformis
SUBPHYLUM HETEROKONTA Thaumatomastix splendida
CLASS BICOSOECIDEA Thaumatomastix type A

12
 2. DIATOMS*
Fiona J. Scott & David P. Thomas

Diatoms are predominantly unicellular organisms, although pseudofilamentous and colonial

aggregations sometimes occur. The cell wall is composed of silica (Si0 2 ) with an organic
coating, the degree of silicification ranging from light to heavy. The resilient nature of the
cell wall has proved an invaluable tool for reconstructing the fossil record. In particular, the
biostratigraphy of fossil diatoms in high latitude oceans has been influential in studies of
past oceanographic circulation and global climatic changes (Gersonde, 1990). The siliceous
walls are variously ornamented with perforations, ribs, spines or minute processes (Fryxell &
Hasle, 1972a), and for almost two centuries the classification of diatoms was based on the
structure and ornamentation of the cell walls.

Diatoms range in size from about 2 to 200 um, and they exhibit in a wide variety of shapes.
They are photosynthetic, generally golden brown, yellow, green or dark brown in colour,
with pigments including chlorophylls a and c, P-carotene, fucoxanthin, diatoxanthin and
diadinoxanthin (Jeffrey et al., 1997). Diatoms are uninucleate and contain chrysolaminarin
and oils as their principal reserve substances. Flagellated cells are only known from male
gametes of centric species, but gliding motility may occur in pennate diatoms possessing a
raphe. Diatoms reproduce vegetatively by binary fission and can form vegetative resting
spores. They can also undergo sexual reproduction during which gametes, auxospores and
'normal' vegetative cells are formed (Hasle & Syvertsen, 1997).

Cells either have concentric or radialsymmetry around a point or points (centric diatoms), or
have a symmetry around a longitudinal axis (pennate diatoms). Round et al. (1990), Medlin
& Priddle (1990) and Hasle & Syvertsen (1997) present excellent accounts of diatom
morphology. Diversity is exceptionally high, including some 285 genera (Round et al., 1990)
and 10,000-12,000 recognised species (Norton et al., 1996). This chapter documents 196
diatom species and infraspecific taxa (115 centric and 81 pennate) from Antarctic waters and
the Southern Ocean.

Historically, diatoms have been classified according to their gross morphology, and a number
of classification schemes have been devised (see discussion in Round et al, 1990). More
recently, molecular techniques have provided an opportunity for a more precise definition of
species and the identification of geographical variants (Graham & Wilcox, 2000); ribosomal
RNA sequences may also prove to be useful in diatom identification (Scholin et al., 1997). In
this treatment we make no assumptions about evolutionary trends, and we list taxa in
alphabetical order within their respective genera and families. The classification followed
here is that of Hasle & Syvertsen (1997) unless indicated otherwise.

Order Centrales
Cells have their valve structure, including the striae, arranged in radial symmetry from a
central point or annulus (cf. Order Pennales, p. 121).

Family Asterolampraceae

Asteromphalus heptactis (Breb.) Ralfs, in Pritchard (1861: 838, pi. 8, fig. 21)

BASIONYM: Spatangidium heptactis Breb. (Brebisson, 1857: 296, pi. 3, fig. 2)

SYNONYMS: Spatangidium ralfsianwn G.Norman (1859: 161, pi. 7, fig. 7)
Asterolampra heptactis (Breb.) Grev. (Greville, 1860: 122), as hepacticus
Asterolampra ralfsiana (G.Norman) Grunow (1876), in Schmidt et al. (1874-1959: pi. 38, figs 5-8)

Class Diatomophyceae or Class Bacillariophyceae.

13
 DIATOMS

Asterolampra areolata A.Mann (1925: 30, figs 5, 6)

ADDITIONAL REFERENCES: Boden (1985: 29), Priddle & Fryxell (1985: 116, figs A, E), Ricard
(1987: figs 54-57), Andreoli et al. (1995: 472, fig. 5)

DESCRIPTION: Cells solitary, discoid. Valves slightly convex, 42-175 um diameter; central
hyaline area occupying c. 25-33% of valve diameter; 7 hyaline rays, one markedly narrower
than the others, extending from centre almost to periphery of valve and ending in a
protruding tubular process. Separating lines within central area bent or branched. Peripheral
sectors areolate; areolae in tangential lines, 5-7 in 10 um, 3 in 10 urn towards the centre.

distribution: Typically found in temperate waters (Hasle & Syvertsen, 1997); Southern
Ocean (Priddle & Fryxell, 1985); Ross Sea (Watanabe, 1982; Andreoli et al., 1995);
Bransfield Strait (Priddle, 1985).

Priddle & Fryxell (1985) noted the difficulty in distinguishing A. heptactis from A. parvulus
(see below), although the latter generally has a smaller valve diameter (22-48 urn).

Asteromphalus hookeri Ehrenb. (Ehrenberg, 1844b: 200, fig. 3)

SYNONYMS: Asteromphalus buchii Ehrenb. (Ehrenberg, 1844b: 200, fig. 4)

Asteromphalus cuvieri Ehrenb. (Ehrenberg, 1844b: 200, fig. 7)

Asteromphalus humboldtii Ehrenb. (Ehrenberg. 1844b: 200, fig. 6)

For further synonyms see Van der Spoel et al. (1973: 529, figs 4, 5).

ADDITIONAL REFERENCES: Ehrenberg (1854: pi. 35A, fig. XXI, 2), Hustedt (1958: 127, figs 88-90),
Kozlova (1962: pi. 9, fig. 6), Priddle & Fryxell (1985: 118), Moisan & Fryxell (1993: 493)

Fig. 2.1a-g

description: Cells solitary, or sometimes in short chains of 2 or 3 cells, discoid. Valves

slightly convex, 25-60 um diameter; central hyaline area occupying c. 33-50% of valve
diameter; 6-9 hyaline rays, one markedly narrower than the others, extending from centre
almost to periphery of valve and ending in a protruding tubular process (Fig. 2.1b); rays
more numerous in larger cells. Separating lines within central area straight. Peripheral
sectors areolate; areolae in tangential lines, 5-9 in 10 um, of uniform size (Fig. 2. Id).

distribution: Prydz Bay, East Antarctica; wide distribution in temperate and subpolar seas
(Hendey, 1937); South Georgia (Hendey, 1937); common throughout Antarctic oceanic areas
(Hargraves, 1968); Indian sector of the Antarctic (Kozlova, 1962); Weddell-Scotia Sea
(Garrisons a/., 1987).

This is one of the most common of the Antarctic Asteromphalus species.

Asteromphalus hyalinus G.Karst. (Karsten, 1905: 90, pi. 8, fig. 15)

ADDITIONAL REFERENCES: Hustedt 1958: 128, figs 84-87),

( Priddle & Fryxell (1985: 120, figs a-f),
Garrison et al. (1987: 566), Moisan & Fryxell (1993: 493)

Fig. 2.2a-d

description: Cells solitary, discoid. Valves slightly convex, 13.5-33 um diameter; central
hyaline area occupying half of valve diameter; 3-5 (-6) hyaline rays, one markedly narrower
than the others, extending from centre almost to periphery of valve and ending in a
protruding tubular process (Fig. 2.2a); rays often slightly curved. Separating lines within the
central area genuflexed and branched. Peripheral sectors areolate; areolae 6-9 between the
hyaline rays, of uniform size (Fig. 2.2b).

distribution: Cape Evans, Ross Sea; Southern Ocean south of Australia; coastal waters of
Vestfold Hills, East Antarctica; Lutzow-Holm Bay (Tanimura et al., 1990); Weddell-Scotia
Sea (Garrison et al., 1987).

14
 DIATOMS

Figure Asteromphalus hookeri. a, LM Valve view of vegetative cell, b, SEM Oblique

2.1.
view, c, Valve view; note straight separating lines, d, SEM Internal valve view of small
SEM
cell, e, SEM
Detail of areolae, external view, f, SEM Detail of areolae, internal view, g,
TEM Shadow cast of valve. Scale bars: a, c = 10 urn; b, f = urn; d, e, g = 5 um.
1

15
 DIATOMS

Figure 2.2. Asteromphalus hyalinus. a, LM Valve with 5 hyaline rays, external view, b,
SEM Oblique view of valve with 6 hyaline rays, c, TEM Valve with 4 hyaline rays and
eroded areolae, d. TEM View through both valves of cell. Scale bars: a-d = 10 urn.

16
 DIATOMS

Some specimens examined for this study were slightly outside the previously recorded cell
diameter range of 15-32 urn (Hasle & Syvertsen, 1997), but in all other respects they agree
with earlier descriptions. A few specimens also had 6 hyaline rays (previous descriptions
reported 3-5). This species is perhaps not as rare in the Antarctic and sub-Antarctic as was
previously thought (Hargraves, 1968).

Asteromphalus parvulus G.Karst. (Karsten, 1905: 90, pi. 8, fig. 14)

ADDITIONAL REFERENCES: Hustedt (1958: 128, fig. 91), Priddle & Fryxell (1985: 116, figs B-D, F);
Hasle & Syvertsen (1997: 139, pi. 23)

Fig. 2.3a-d

description: Cells solitary, discoid. Valves slightly convex, 22-48 pm diameter; central
hyaline area occupying c. 50-75% of valve diameter; 6 or 7 hyaline rays, one markedly
narrower than the others, extending from centre almost to periphery of valve and ending in a
small tubular process. Separating lines within the central area broken or with small
branchlets in the middle. Peripheral sectors areolate; areolae rather coarse, of uniform size,
8-10 in 10 um, in tangential lines. Chloroplasts spherical, numerous.

DISTRIBUTION: Southern Ocean, offshore MacRobertson Land, 65°38'S 68°07'E; coastal

waters near Davis Station, East Antarctica; Franklin I., Ross Sea; South Orkney Is. (Ferrario
& Ferreyra, 1987); Bellingshausen Sea, Drake Passage (Hendey, 1937); Indian sector of the
Antarctic Ocean (Kopczyhska et al., 1986); Weddell-Scotia Sea (Garrison et al, 1987).

Asteromphalus roperianus (Grev.) Ralfs, in Pritchard (1861: 838)

BASIONYM: Asterolampra roperiana Grev. (Greville, 1860: 120, pi. 4, fig. 14)

SYNONYM: Asterolampra roperiana Castrac. (Castracane, 1886: 133, pi. 5, fig. 3), nom. Meg.

ADDITIONAL REFERENCES: Priddle & Fryxell (1985: 122, figs A, B), Hasle & Syvertsen (1997:
139, pi. 24)

description: [after Hasle &

Syvertsen (1997)] Cells solitary, discoid. Valves 80-120 urn
diameter; central hyaline area occupying c. one-third of valve diameter; 7 hyaline rays, one
markedly narrower than the others, extending from centre almost to periphery of valve.
Separating lines within the central area obscurely bent or with small branchlets in the middle.
Peripheral sectors areolate; areolae 9-1 1 in 10 pm.

distribution: Southern Ocean (Priddle & Fryxell, 1985); Wilkes Land, East Antarctica
(Bunt & Wood, 1963).

This is regarded as a predominantly warm-water species (Simonsen, 1974). Although not

seen by us, it is included here due to having been reported from the Southern Ocean and
Wilkes Land.

Eucampia antarctica (Castrac.) Mangin (1914: 480, figs 7, 8) var. antarctica

BASIONYM: Moelleria antarctica Castrac. (Castracane, 1886: 98, pi. 18, fig. 8)

SYNONYMS: Eucampia balaustium Castrac. (Castracane, 1886: 97, pi. 18, fig. 5), Hendey (1937: 285),
Manguin (1954: 19), Hustedt (1958: 136, figs 40-43), Jouse et al. (1962: 82, pi. 5, figs 14-17), Hoban et al.
(1980: 592, figs 1-14), Syvertsen & Hasle (1983: 181, figs 4, 84-129)
Eucampia balaustium Castrac. var. minor Castrac. (Castracane, 1886: 98, pi. 18, fig. 6)

Eucampia antarctica (Castrac.) Mangin f. spiralis Mangin (1915: 63)

Eucampia balaustium Castrac. f. hiberna Heiden, in Heiden & Kolbe (1928: 535)
Eucampia balaustium Castrac. f. aestiva Heiden, in Heiden & Kolbe (1928: 536)
ADDITIONAL REFERENCES: Priddle & Fryxell (1985: 66, figs A-D), Fryxell et al. (1989: 638),
Hasle & Syvertsen (1990c: 129, pi. 16.1, figs 7-13), Fryxell & Prasad (1990: 33), Moisan &
Fryxell (1993:493)

Fig. 2.4a-f

17
 DIATOMS

description: Cells solitary, in doublets or spiralling chains, the spiralling a result of

asymmetrical growth of the valve and girdle bands; occurring as vegetative cells (Fig. 2.4a) or
heavily silicified 'winter' resting spores (Fig. 2.4b). Vegetative cell pervalvar axis 70-120 urn;
resting spore pervalvar axis 30-52 urn; ratio between apical and pervalvar axes variable.
Horns broad, flattened and ocellate (vegetative cell, Fig. 2.4a) or tapering (secondary valve
of resting spore, Fig. 2.4b). Valves elliptical; apical axis 18-92 urn; rows of large areolae,
3-10 in 10 um, radiating from a central annulus (Fig. 2.4c-e). Labiate process solitary,
marginal (Fig. 2.4c). Vela are complex cribra (Fig. 2.4e, f). See Syvertsen & Hasle (1983)
for a full description (as E. balaustium).

distribution: Prydz Bay, East Antarctica; type locality (of E. balaustium) is recorded as
Antarctic Ocean (Castracane, 1886); Weddell Sea (Buck & Garrison, 1983; Fryxell, 1989);
Heard I. (Manguin, 1954); Wilkes Land, Antarctica (Bunt & Wood, 1963); Indian Ocean
sector of the Antarctic (Kopczyhska et al., 1986); Weddell-Scotia Sea (Garrison et al., 1987);
coastal waters off Syowa Station, East Antarctica (Ishikawa et al., 2001).

Eucampia antarctica (Castrac.) Mangin var. recta (Mangin) G. A. Fryxell & A. K.S.Prasad
(1990: 34, figs 2-19)
BASIONYM: Eucampia antarctica (Castrac.) Mangin f. recta Mangin (1915: 63, figs 41-44)

SYNONYM: Hemiaulus antarcticus Ehrenb. (Ehrenberg, 1844b: 203, fig. 8)

Fig. 2.5a, b

description: Cells solitary, in doublets or straight to slightly curved chains, occurring as

vegetative cells or heavily silicified 'winter' resting spores. Valves elliptical; apical axis 39-
116 um; areolae 7-8 in 10 um. Horns of vegetative cells broad, flattened and ocellate (Fig.
2.5a). Labiate process solitary, c. 2 areolae from the margin. Vela are complex cribra (Fig.
2.5b). See Fryxell & Prasad (1990) for a full description.

distribution: Franklin L, 76°09'S 168°20.366'E, Ross Sea; Weddell Sea, 72°20.1'S and
74°39.12'S (Fryxell & Prasad, 1990).

Eucampia antarctica var. recta is distinguished from the type variety principally in having
only a slight curvature of the chains (in narrow girdle view only). Moreover, var. recta has
broader valve dimensions (39-1 16 um) and is of finer structure (areolae 7-8 in 10 um).

The distribution of these two varieties overlap in the Weddell Sea, but while var. antarctica
has a polar to subpolar distribution with a northernmost record at c. 56°S (Manguin, 1954),
var. recta appears to be an Antarctic endemic, with a northernmost record at 64°48'S
(Fryxell & Prasad, 1990).

Trigonium arcticum (Brightw.) Cleve (1868: 663)

BASIONYM: Triceratium arcticum Brightw. (Brightwell, 1853: 250, pi. 4, fig. 1
1)

synonym: Biddulphia arctica (Brightw.) CS.Boyer (1901 714) :

ADDITIONAL REFERENCES: Hendey (1937: 282, pi. 10, fig. 1), Hoban (1983: 277, figs 37-42).
Everitt & Thomas (1986: 9, fig. 3B), Round et al. (1990: 254)

Fig. 2.6a-f

DESCRIPTION: Cells solitary or in zig-zag chains, triangular in valve view (Fig. 2.6a-c) and
slightly elevated at the corners, rectangular in girdle view. Apical axis 150-350 um; areolae
polygonal, radiating in rows from the centre, 3-4 per 10 um in central region, smaller at valve
corners where they form pseudocelli; central grouping of small labiate processes (Fig. 2.6f);
valves with external cribrum-like rotae (Fig. 2.6b, e) and internal foramina (Fig. 2.6c, 0'
Girdle short, covered in fine punctae arranged in straight vertical lines (Fig. 2.6a).

distribution: Inexpressible I., 74°54'S 163°46'E, Ross Sea; inshore Davis Station, Prydz
Bay, East Antarctica (Everitt & Thomas, 1986); Southern Ocean, 62°59'S 57°28'W, off
Enderby Land (Hendey, 1937); Wilkes Land, Antarctica (Bunt & Wood, 1963).
 DIATOMS

Figure 2.3. Asteromphalus parvulus. a, LM Valve with 7 hyaline rays, b, LM Valve with 7
hyaline rays, c, SEM External valve view, d, SEM Internal valve view. Scale bars: a-d = 10 urn.

19
 DIATOMS

***** ^'^ :5*i

Figure 2.4. Eucampia antarctica var. antarctica. a, LM First step in semi-endogenous spore-
formation from a vegetative cell, b, SEM Resting spore, epitheca towards left, secondary
valve with tapered horns on right, c, SEM Vegetative cell valve; note ocelli and labiate
process, d, SEM Resting spore valve, e, SEM Vegetative cell valve; note ridged ocelli, f, SEM
Vegetative cell valve cribra. Scale bars: a, b = 10 urn; c = 1 um; d, e = 5 urn; f = 500 nm.

20
 )

DIATOMS

Figure 2.5. Eucampia antarctica var. recta, a, SEM Valve of vegetative cell, b, SEM
Detail of valve areolae. Scale bars: a = 10 urn; b = 5 urn.

Everitt & Thomas (1986) reported this as a benthic diatom that can become mixed into the
water column in winter. Hendey (1937) described its habitat as and commonly
littoral
epiphytic.

Family Chaetoceraceae
This family is represented by 25 species of the genus Chaetoceros. Gran (1897) recognised
two subgenera: Phaeoceros, large, robust forms with numerous, small chloroplasts and thick
setae armed with spines (later corrected to subgenus Chaetoceros by Hernandez-Becerril,
1993); and Hyalochaetae, small, delicate forms with usually one or a few plate-like
chloroplasts and thin setae.

Chaetoceros adelianus Manguin (1957: 1 18, pi. 3, fig. 16a-f)

ADDITIONAL REFERENCES: Manguin (1960: 274, pi. 7, figs 83-88), Priddle & Fryxell (1985: 20,
figs on p. 2 1

Fig. 2.7a

description: Cells chain-forming, heavily silicified, rectangular in girdle view; pervalvar

axis 42-48 urn. Valves oval-elliptical; apical axis 37-43 urn, depressed in the centre and
with a small central spine. Apertures between cells small. Setae long, coarse, orientated
almost perpendicular to pervalvar axis, spinose; in valve view, one seta ±continues in the
direction of the (longer) apical axis, while the other curves away from it (Manguin, 1957:
fig. 16c).

distribution: Southern Ocean, south of Australia; Terre Adelie (Manguin, 1957, 1960).

Chaetoceros aequatorialis Cleve var. antarcticus Manguin (1960: 274, pi. 8, fig. 89)
SYNONYM: IChaetoceros pendulus G.Karst. (Karsten, 1905: 1 18, pi. 15, fig. 7) (see Hasle & Syvertsen, 1997)

21
 DIATOMS

ADDITIONAL REFERENCES: Priddle & Fryxell (1985: 22, fig. on p. 23), Hasle & Syvertsen (1997:
193, pi. 38, tab. 46)

Fig. 2.8a-d

description: Cells solitary, heterovalvate, rectangular in girdle view; pervalvar axis 6.5-9 urn.
Valves elliptical; apical axis 10-11 urn, with a small off-centre tubular extension of the
process. Setae long, coarse, spinose; setae on the upper valve slightly enlarged at the point of
emergence from inside the valve margin, curving downwards and becoming convergent
towards their ends; setae from lower valve ±parallel to pervalvar axis.

distribution: Coastal waters near Davis Station, East Antarctica; type locality, Terre
Adelie, 63°25'S 143°09'E (Manguin, 1960).

The type variety is regarded as a warm-water diatom, Hendey (1937) recording it as "an
oceanic species, widely spread throughout the Indian Ocean". The only morphological
differences between the var. aequatorialis and var. antarcticus seem to be the size (apical
axes c. 25 um and 10-11 um, respectively) and the orientation of the setae that arise from
the upper valve; those of the type variety remain parallel for their entire length, while those
of var. antarcticus converge distally. Chaetoceros pendulus has a similar morphology to
C. aequatorialis var. antarcticus, but it has smooth rather than spinose setae.

Chaetoceros atlanticus Cleve (1873: 1 1, pi. 2, fig. 8)

SYNONYMS: Chaetoceros dispar Castrac. (Castracane, 1886: 76, pi. 8, fig. 6)

Chaetoceros polygonum Schutt (1895: 46, pi. 5, fig. 24)

Chaetoceros audax Schutt (1895: 47, pi. 5, fig. 25)
See VanLandingham (1968: 701 ) for full synonymy.

ADDITIONAL REFERENCES: Karsten (1905: 115, pi. 15, fig. 9), Hustedt (1926: 641, figs 363, 364),
Hendey (1937: 290), Manguin (1954: 15), Hustedt (1958: 135), Evensen & Hasle (1975: 157, figs
6-11), Moisan& Fryxell (1993: 493), Hasle & Syvertsen (1997: 196, pi. 39a)

Fig. 2.9a-c

DESCRIPTION: Cells united to form rectangular in girdle view; pervalvar

stiff straight chains,
axis 12-40 um; girdle distinctly Valves oblong-elliptical; apical axis 10-46 um;
visible.
surface flat, margins curved, witha small central spine. Apertures large, although varying in
size, somewhat rectangular. Setae strong, to 200 um long, arising from the valve surface
slightly within the margin, usually stiff and straight, but sometimes slightly inflated and
markedly sigmoidal with apices bent inwards (Fig. 2.9b, c); setae bearing lines of fine
punctae and small spines. Chloroplasts numerous, small, spherical, distributed into the setae.

distribution: South Atlantic, 61°32'S 0°00' to 51°29'S 0°08'W (Hustedt, 1958); Southern
Ocean (Hendey, 1937; Priddle & Fryxell, 1985); north polar and temperate waters (Hendey,
1937); Ross Sea (Andreoli et al., 1995); Weddell-Scotia Sea (Garrison et al, 1987);
KERFIX, 50°40'S 68°25'E, Southern Ocean (Kopczyhska et al., 1998).
The long setae of Chaetoceros atlanticus distinguish it from C. bulbosus, the setae of which
are markedly inflated and much shorter (c. 20 um).

Chaetoceros brevis Schutt (1895: 38, fig. 4)

ADDITIONAL REFERENCES: Cupp (1943: 129, fig. 82), VanLandingham (1968: 705), Hasle &
Syvertsen (1997: 210)

Fig. 2.14a

description: Cells in short chains; apical axis 8-17 um. Setae almost straight. Chloroplast
single.

distribution: Weddell Sea (Garrison et a/., 1987, as C. breve); Ross Sea (Watanabe, 1982);
Scotia Sea (Garrison & Buck, 1989b).

22
r^^ DIATOMS

Figure 2.6. Trigonium arcticum.a, SEM Vegetative cell, b, SEM Valve, external surface;
note radiating areolae, c, SEMValve, internal surface, d, SEM Small areolae forming a
pseudocellus on external surface of valve, e, SEM Cribra on valve external surface, f, SEM
Cribra on valve internal surface. Scale bars: a-c = 100 um; d-f = 10 urn.

23
 DIATOMS

Figure 2.7. Chaetoceros species, a, C. adelianus, LM Short chain of cells, b, C. castracanei,

SEM terminal and adjacent cells of a chain; note central protuberances bisecting the aperture.
c, C. curvatus, SEM Solitary cell in girdle view, d, C. gaussii, SEM Solitary cell in girdle
view. Scale bars: a-d = 10 jam.

24
 DIATOMS

Figure 2.8. Chaetoceros aequatoralis var. antarctica. a, LM Cell in girdle view, b, SEM
Collapsed cell with clear view of central processes, c, SEM View of upper valve with emergent
setae and off-central processes, d, SEM View of lower valve. Scale bars: a-d = 10 um.

25
 DIATOMS

Figure 2.9. Chaetoceros species, a-c, LM Upper valve and setae, b, SEM
C. atlanticus. a,
Vegetative cell; note long central spine, c, SEM
Vegetative cell with sigmoidal setae, d-f, C.
bulbosus. d, SEM Valve, internal view of valve, base of setae and central labiate process, e.
LM Vegetative cell in girdle view; note striated setae, f, SEM Vegetative cell; setae
appearing fenestrated. Scale bars: a, b, e = 10 um; c, d, f = 5 urn.

26
 DIATOMS

According to Hasle & Syvertsen (1997) "there seems to be a general consensus that positive
identification of C. brevis is problematic".

Chaetoceros bulbosus (Ehrenb.) Heiden, in Heiden & Kolbe (1928: 526, pi. 9, fig. 171)

BASIONYM: Dicladia bulbosa Ehrenb. (Ehrenberg, 1844: 20; 1854, pi. 35A/21, fig. 10)

SYNONYMS: Chaetoceros radiculum Castrac. (Castracane, 1886: 79, text fig.)

? Chaetoceros schimperianus G.Karst. (Karsten, 1905: 1 17, pi. 15, fig. 2; pi. 16, fig. 4)

IChaetoceros bulbosus G.Karst. f. schimperiana (G.Karst.) Heiden, in Heiden & Kolbe (1928: 528, pi. 10,
fig. 174; pi. 11, fig. 176), as schimperana
Chaetoceros atlanticum f. bulbosum (Ehrenb.) Hargraves ( 1968: 26, figs 28-31)

ADDITIONAL REFERENCES: Hustedt (1958: 135), Moisan & Fryxell (1993: 493), Andreoli et al
(1995: fig. 8)

Fig. 2.9d-f

description: Cells solitary or in short chains, somewhat octagonal in girdle view; pervalvar
axis 10-25 urn. Valves elliptical; apical axis 40-50 urn; surface flat, sloping towards the
margins, with the centre slightly raised and with a conspicuous central process. Apertures
large, although varying in size, somewhat rectangular. Setae strong, c. 20 urn long, arising
from the valve surface slightly inside the margin, bulbous at their base, tapering distally (Fig.
2.9d); setae often striate, bearing lines of fine punctae. Chloroplasts small, spherical,
numerous, distributed into the setae.

distribution: Coastal waters near Davis Station, East Antarctica; Weddell Sea (Buck &
Garrison, 1983; Moison & Fryxell, 1993); Ross Sea (Watanabe, 1982; Andreoli et al., 1995)
Weddell-Scotia Sea (Garrison et al., 1987); Liitzow-Holm Bay (Tanimura et al., 1990)
Prydz Bay, East Antarctica (Boden, 1985); Wilkes Land, Antarctica (Bunt & Wood, 1963)
between the South Shetlands and Cape Horn (Hendey, 1937, as C. radiculum); Southern
Ocean, between 51° and 49°S, 0°08' and 0°07'W, and 53°43'S 0°20'W, 49°S 0°07'W and
46°32'S 0°02'W (Hustedt, 1958); Southern Ocean-Davis Sea, 47-65°S (Heiden & Kolbe,
1928); KERFIX, 50°40'S 68°25'E, Southern Ocean (Kopczyhska et al., 1998).

Chaetoceros bulbosus was referred to by Priddle &

Fryxell (1985) as "C bulbosus complex",
including the atlanticus, bulbosus, schimperianus and cruciata phases. On morphological
grounds, Hargraves (1968) suggested that C. bulbosus was a cold-water form of
C. altlanticus, but he noted that the "type form also occurred at same stations as theform
bulbosum".

Chaetoceros castracanei G.Karst. (Karsten, 1905: 1 16, pi. 15, fig. 1)

ADDITIONAL REFERENCES: Castracane (1886: 80, text fig., as "Chaetoceros sp. nov.?"), Manguin
(1960: 278, pi. 8, figs 92, 93; pi. 27, fig. 327), Priddle & Fryxell (1985: 26), Hasle & Syvertsen
(1997: 199, pi. 39)

Fig. 2.7b

DESCRIPTION: Cells in straight chains, isovalvate, rectangular in girdle view; pervalvar axis
11-17 um. Valves elliptical, 10.5-23 urn diameter, with a central protuberance. Apertures
between cells narrow. Setae long, thin, finely spinose for much of their length, arising from
the valve surface slightly inside the margin, those of adjacent cells at first fused, then lying
perpendicular to the chain axis. Chloroplasts small, spherical.

distribution: Coastal waters off Davis Station, East Antarctica; Terre Adelie (Manguin,
1960); Weddell Sea (Garrison & Buck, 1989b); Bransfield Strait (Priddle, 1985); Southern
Ocean (Priddle & Fryxell, 1985); Weddell-Scotia Sea (Garrison et al., 1987); Ross Sea
(Watanabe, 1982).

27
 DIATOMS

Chaetoceros concavicornis Mangin (1917: 9, figs 5-7)

ADDITIONAL REFERENCES: VanLandingham (1968: 709), Evensen & Hasle (1975: 158, figs 15-22),
Hasle & Syvertsen (1997: 199, pi. 40)

Fig. 2.14b

description: Cells in straight chains, heterovalvate, rectangular in girdle view, with upper
face rounded; pervalvar axis 13-20 urn. Valves elliptical, 12-30 um diameter; upper valve
rounded, with setae arising from near the centre; lower valve flat, with setae arising from
inside valve margin; small central spine present. Apertures between cells small, ±trapezoid.
Setae long, thin at point of emergence from valve, becoming broader distally, finely spinose
for much of their length. All setae curving back towards lower end of chain. Chloroplasts
small, spherical.

distribution: Bransfield Strait (Kopczyhska & Ligowski, 1985); Ross Sea (Watanabe,
1982).

Hasle & Syvertsen (1997) reported this as a "cosmopolitan species, common in temperate
and northern cold waters but absent from southern cold waters". It is included here because
of the few records from more southerly locations. Its morphology and size are similar to
C. peruvianus, cells of which are also heterovalvate and with downward-curving, spinose
setae. However, C. peruvianus has a pronounced constriction between the valve mantles and
a narrow girdle in girdle view (this area is straight in C. concavicornis). Moreover, the setae
arise as a distinct, vault-like structure from the centre of the upper valve.

Chaetoceros convolutus Castrac. (Castracane, 1886: 78, text fig.)

SYNONYMS: Chaetoceros criophilus auct. non Castracane (1886), sensu Cleve (1897: 20, pi. 1, fig. 6)

Chaetoceros brightwellii auct. non Cleve (1873), sensu Gran (1897: 1 1, pi. 1, fig. 1)

additional references: Meunier (1910: 218, pi. 24, figs 17-19), Hustedt (1930: 668, fig. 378),
Lebour (1930: 119, fig. 86), Hendey (1937: 293), Manguin (1960: 279, pi. 8, figs 94-97), Moisan &
Fryxell (1993: 493), Priddle & Fryxell (1985: 28, figs A-F), Hasle & Syvertsen (1997: 199, pi. 41)

Fig. 2.10a, b

description: Cells united to form curved or slightly twisted chains, heterovalvate; pervalvar
axis 15-28 um. Valves circular; apical axis 10-30 um; upper valve domed; lower valve flat;
small central spine often obscured. Apertures between cells small, frequently obscured. Setae
long, thin, bent towards lower end of chain, spinose throughout their length; setae of upper
valve arising from the centre, forming a vault-like structure, those of lower valve arising
from near the valve margin; setae of lower valve of terminal cell often converge inwards to
end of chain and cross over. Chloroplasts numerous, small, spherical.

distribution: Southern Ocean, south of Australia; Franklin I., 76°09.59'S 168°20.36'E,

Ross Sea; Weddell Sea (Moisan & Fryxell, 1993); Weddell-Scotia Sea (Garrison et al,
1987); Southern Ocean (south-west of Kerguelen Is.; Hendey, 1937); Atlantic and northern
European seas (Ricard, 1987); Terre Adelie (Manguin, 1960).

Chaetoceros criophilus Castrac. (Castracane, 1886: 78, text fig.)

ADDITIONAL REFERENCES: Hendey (1937: 295, pi. 13, fig. 7), Hustedt (1958: 135), VanLand-
ingham (1968: 712), Priddle & Fryxell (1985: 30, as C. criophilum), Everitt & Thomas (1986: 5,

fig. 2A), Fryxell (1989: 10, figs 33, 34), Hasle & Syvertsen (1997: 193, pi. 38)

Fig. 2.10c, d

description: Cells solitary or chain-forming, ±rectangular in girdle view, heterovalvate;

pervalvar axis 15-55 um. Valves almost circular; apical axis 16-50 um diameter; upper
valve convex; lower valve flat. Setae long, curved back towards lower end of chain, spinose
throughout their length; setae of upper valve arising near the centre as part of the valve

28
 DIATOMS

Figure 2.10. Chaetoceros species, convolutus. a, LM Chain of cells, b, SEM Cell

a, b, C. in
girdle view; note vault-like structure formed by setae arising from upper valve, c, d, C.
criophilus. c, LM, Chain of cells, some setae broken, d, SEM, Adjoining vegetative cells;
note spinose setae arising as part of the upper valve surface, e, f, C. dichaeta. e, LM
One and
a half vegetative cells; setae broken, f, SEM Internal view of valve with bases of setae and
central spine. Scale bars: a-e = 10 um; f = 1 urn.

29
 DIATOMS

Figure 2.11. Chaetoceros dichaeta. a. LM Solitary cell in girdle view. b. SEM Short chain
of cells, c. SEM Short chain of cells, d, SEM Detail of valve with setae and central spine.
Scale bars: a-d = 10 (am.

30
 DIATOMS

/surface (Fig. 2. 1 Od), those of lower valve arising from near valve margin; a characteristic
notch between setae and valve face on the lower valve. Chloroplasts numerous, small,
spherical, sometimes distributed into the setae.

distribution: Inshore waters of Davis Station, East Antarctica (Everitt & Thomas, 1986);
South Georgia, Weddell Sea, Bellingshausen Sea, Ross Sea (Hendey, 1937); Lutzow-Holm Bay
(Tanimura et al., 1990); widespread in Antarctic and South Atlantic especially from 61°32'S
0°00' to 53°43'S 0°20'W (Hustedt, 1958); Weddell-Scotia Sea (Garrison et al, 1987).

This is a characteristic Antarctic diatom (Hendey, 1937). Although resting spores are not
known, lipid-rich, relatively dormant cells may be the 'winter' forms (Fryxell, 1989).

Chaetoceros curvatus Castrac. (Castracane, 1886: 77, text fig.)

ADDITIONAL REFERENCES: Mangin (1915: 36, figs 15, 16), Hendey (1937: 294), Hustedt (1958:
135)

Fig. 2.7c

description: Cells usually solitary, short chains of 2-6 cells,

sometimes united in
±rectangular in 10-15 um. Valves elliptical to
girdle view, heterovalvate; pervalvar axis
circular; apical axis 15-36 um diameter; upper valve convex; lower valve flat or concave.
Setae emerging from centre of valves, those of upper valve distinctly separate and abruptly
reflexed, those of lower valve divergent and lying ±parallel to upper setae. Chloroplasts
small, numerous.

DISTRIBUTION: Southern Ocean, south of Australia; South Atlantic at 51°29'S 0°08'VV

(Hustedt, 1958); Petermann I., Bransfield Strait (Mangin, 1915); temperate South Atlantic

and around the coasts of South Africa (Hendey, 1937).

This species resembles Chaetoceros criophilus, but the setae emerging from the upper valve
are separate rather than fused, as in C. criophilus (Fig. 2.10d).

Chaetoceros curvisetus Cleve (1889: 55)

ADDITIONAL REFERENCES: Cleve (1894: 12, pi. 1, fig. 5), Hustedt (1958: 135), VanLandingham
(1968: 713), Evensen &
Hasle (1975, figs 25, 26), Priddle & Fryxell (1985: 44, figs A-C), Rines
&Hargraves(1988: 71, figs 141, 142), Hasle & Syvertsen (1997: 21 1, pi. 44, tab. 51)

Fig. 2.14c

description: Cells in spirally curved chains, ±rectangular in broad girdle view, isovalvate;
pervalvar axis 20-38 um; adjacent cells connected by the raised corners. Valves elliptical,
concave; apical axis 7-30 um diameter. Apertures between adjacent cells rhombic, oval or
circular. Central labiate process present onlyon terminal cell of chain, short, flattened, with
no protrusion or thickening on the inside (Evensen & Hasle, 1975). Setae with characteristic
curvature, all directed toward the outside of the spiral, thin, tapering distally. Chloroplast
single. Resting spores smooth.

DISTRIBUTION: Ross Sea (Watanabe, 1982); Weddell Sea (Garrison & Buck, 1989b);
cosmopolitan, mostly temperate and warm waters (Hasle & Syvertsen, 1997); Antarctic
sector of the Atlantic Ocean, 64°10'S O^'W, 59°S 0°05'W (Hustedt, 1958); Southern
Ocean (Priddle & Fryxell, 1985).

Chaetoceros debilis Cleve (1894: 13, pi. 1, fig. 2)

ADDITIONAL REFERENCES: VanLandingham (1968: 714), Garrison (1991b: 22), Hasle & Syvertsen
(1997: 211, pi. 44, tab. 51)

Fig. 2.14d

description: Cells chain-forming in distinct spirals, rectangular in girdle view, with valves
of adjacent cells not touching and apertures rectangular, sometimes slightly constricted in the
centre. Valves flat to slightly convex; apical axis 8-40 um. Setae long, thin, smooth,

31
 DIATOMS

crossing slightly after emergence from the valves, extending outwards from the spiral.
Chloroplast single.

distribution: Off southern California (Cupp, 1943); Antarctic (Garrison, 1991b);

cosmopolitan, mainly cooler waters (Hasle & Syvertsen, 1997).

Chaetoceros debilis appears to be rare in the Antarctic. It is similar to C. curvisetus, but the
valves of C. debilis are convex and the cells do not touch at the corners as
flat to slightly

they do in C. curvisetus. The cell chains of C. debilis are markedly spiralled, not simply
curved.

Chaetoceros dichaeta Ehrenb. (Ehrenberg, 1844b: 200)

SYNONYMS: Chaetoceros distans Cleve (1873: 9, fig. 11a, b)

Chaetoceros remotus Cleve & Grunow (1880: 120)

Chaetoceros janischianum Castrac. (Castracane, 1886: 77, text fig.)

additional REFERENCES: Hustedt (1927-30: 648, fig. 367), Hendey (1937: 291, pi. 6, figs 9, 10),
Manguin (1954: 15), Hustedt (1958: 136), Manguin (1960: 282, pi. 28, figs 330-332),
VanLandingham (1968: 718), Evensen & Hasle (1975: 157, figs 1-5), Priddle & Fryxell (1985:
32), Moisan & Fryxell (1993: 493), Andreoli et al. (1995: 472, fig. 8), Hasle & Syvertsen (1997:
196, pi. 39, tab. 47)

Figs2.10e, f;2.11a-d

description: Cells chain-forming, often united by parent girdle, narrow in girdle view,
isovalvate; pervalvar axis 10^0 um. Valves elliptical to circular; apical axis 7-50 urn; valve
face flat, with a central spine. Apertures between cells large, polygonal. Setae arising well
inside the valve margin, running parallel to pervalvar axis for c. 1 cell length, then bending
outwards almost perpendicular to chain axis; terminal setae bending towards the chain axis.
Chloroplasts numerous, small, spherical or discoid, distributed into the setae.

distribution: Southern Ocean, south of Australia; Drygalski Glacier, 75°29'S 165°19'E,

Ross Sea; Southern Ocean, offshore Wilkes Land, 63°50'S 104°37'E; widespread in
Antarctic waters and temperate zones of the South Atlantic (Hustedt, 1958; Hasle &
Syvertsen, 1997); Indian Ocean sector of the Antarctic (Kopczyhska et al, 1986); Weddell-
Scotia Sea (Garrison et al., 1987); Terre Adelie (Manguin, 1960).

Chaetoceros flexuosus Mangin (1915: 45, fig. 27)

additional REFERENCES: Priddle & Fryxell (1985: 34, figs A-D), Hasle & Syvertsen (1997: 203,
pi. 41)

Fig. 2.12a, b

description: Cells chain-forming, attached by setae of adjacent cells twisted around one
another in 1 or 2 turns, heterovalvate, rectangular in girdle view; pervalvar axis 9-20 um.
Valves elliptical to circular; apical axis 9-15 um; upper valve ±flat; lower valve slightly
concave. Chloroplasts numerous, small, spherical or discoid, distributed into the setae
(Fryxell & Medlin, 1981).

distribution: Coastal waters off Davis Station, East Antarctica; Roosen Channel, Peltier
Channel, Matha Bay, Petermann I., Antarctic Peninsula (Mangin, 1915); southern cold-water
regions (Priddle & Fryxell, 1985; Hasle & Syvertsen, 1997); Weddell-Scotia Sea (Garrison et
al., 1987); Drake Passage (Garrison et al., 1987); Indian Ocean sector of the Antarctic

(Kopczyhska et al., 1986).

Chaetoceros gaussii Heiden & Kolbe (1928: 534, pi. 12, fig. 181)

ADDITIONAL REFERENCE: Manguin (1960: 285, pi. 10, fig. 105; pi. 28, fig. 335)

Fig. 2.7d

32
 DIATOMS

Figure 2.12. Chaetoceros species, a, b, C. flexuosus. a, SEM Broken chain of cells; note
characteristic twisting of setae, b, SEM Recently divided cells retained within parent
copulae. c, d, C. hendeyi. c, LM Short chain of cells in girdle view, d, LM Cells in girdle and
apical view. Scale bars: a, b = 10 p.m; c, d = 20 p.m.

33
 DIATOMS

Figure 2.13. Chaetoceros species, a, C. neglectus, SEM Chain of cells, b, C. peruvianus,

LM Solitary cell, c, C. radicans, SEM Chain of cells with characteristically spinose setae, d,
C. simplex, SEM Solitary cell. Scale bars: a. c = 5 urn; b, d = 10 urn.

34
 DIATOMS

description: Cells solitary, rarely remaining attached after cell division, isovalvate,
rectangular in girdle view; pervalvar axis 28-88 urn. Valves elliptical, ±concave; apical axis
47-78 pm. Setae arising from narrow ends of valve margins, at first diverging, then lying in
the transapical plane.

distribution: Southern Ocean, south of Australia; type locality, "Gaussberg", Davis Sea,
"Gauss" Expedition of 1901-02 (Heiden & Kolbe, 1928); Terre Adelie, 66°47'S 141°25'E
(Manguin, 1960).

The single specimen collected in the present study measured 70 x 70 urn; those reported by
Manguin were considerably smaller (apical axis 47-58 urn; pervalvar axis 28-42 urn).

Chaetoceros hendeyi Manguin (1960: 285, pi. 10, figs 106-109)

ADDITIONAL REFERENCES: Priddle & Fryxell (1985: 36, figs A-D), Hasle & Syvertsen (1997: 203)

Fig. 2.12c, d

description: Cells in short straight chains, octagonal in girdle view, isovalvate; apical axis
14-27 urn; pervalvar axis 11.5-16 pm. Apertures between adjacent cells large, hexagonal.
Setae smooth, slightly inflated towards the base, ±at right angles to the pervalvar axis.
Chloroplasts small, spherical, numerous, distributed throughout the cell and into the setae.

distribution: Weddell-Scotia Sea (Garrison et al., 1987); southern cold-water regions

(Priddle & Fryxell, 1985); Terre Adelie (Manguin, 1960).

Chaetoceros hendeyi is supposedly endemic to southern cold-water regions, but it has rarely
been reported since the original description. Some cells found in the present study were
larger (apical axis 17-27 urn) than previously documented (14-18 pm). Hasle & Syvertsen
(1997: 203) noted the possibility that this might be conspecific with C. borealis Bailey from
cold-water regions of the Northern Hemisphere.

Chaetoceros laciniosus Schiitt (1895: 38, pi. 4, fig. 5)

ADDITIONAL REFERENCES: Hendey (1937: 301), Bunt & Wood (1968), VanLandingham (1968:
728), Boden (1985), Hasle & Syvertsen (1997: 209, pi. 43, tab. 50)

Fig. 2.14e

DESCRIPTION: Cells in straight loose chains, isovalvate; apical axis 9-42 pm. Apertures
between adjacent cells square to rectangular. Setae thin, at first parallel to pervalvar axis,
then abruptly diverging almost perpendicularly.

distribution: Ross Sea (Watanabe, 1982); Prydz Bay, East Antarctica (Boden, 1985); South
Atlantic, 49°58.75'S30°10'W (Hendey, 1937).
Although the distribution of C. laciniosus was reported by Hasle & Syvertsen (1997) as
"northern cold water region to temperate?", there are a few earlier reports from Antarctic
waters.

Chaetoceros neglectus G.Karst. (Karsten, 1905: 1 19, pi. 16, fig. 5)

ADDITIONAL REFERENCES: Hendey (1937: 303), Hustedt (1958: 136), Manguin (1960: 286, pi. 11,

figs 112, 113; pi. 28, fig. 336), Priddle & Fryxell (1985: 46, figs A-C), Hasle & Syversten (1996:
207, pi. 44)

Fig. 2.13a

description: Cells chain-forming, weakly silicified, rectangular in girdle view, isovalvate;

pervalvar axis 5.5-10 urn. Valves flat, elliptical; apical axis 8.5-15 pm, no central spine
visible. Apertures between cells large, hexagonal. Setae long, thin, arising from narrow ends
of valve margins, diverging and lying perpendicular to the flexuous cell chain. Chloroplast
single, parietal.

35
 DIATOMS

distribution: Southern Ocean, south of Australia; Weddell-Scotia Sea (Garrison et al,

1987); southern cold-water region (Hasle & Syvertsen, 1997); Ross Sea (Watanabe, 1982);
Terre Adelie (Manguin, 1960); South Atlantic, Antarctic sector, between 51°32'S 0°08'W
and 41°30'S 0°03'W (Hustedt, 1958); South Georgia, South Sandwich Group, South
Shetlands, Weddell Sea, Bellingshausen Sea (Hendey, 1937).

Cells collected from the Southern Ocean were slightly smaller (apical axis 5.8-6.4 urn) than
previous records. Chaetoceros neglectus and C
socialis are similar small species, the latter
characterised by having 3 short and 1 long setae per cell. Chaetoceros neglectus is also
morphologically similar to C. dichaeta, but the latter has a long central spine on each valve
and numerous small chloroplasts distributed into the setae.

Chaetoceros neogracilis (Schiitt) VanLand. (VanLandingham, 1968: 733)

BASIONYM: Chaetoceros gracile Schiitt (1895: 42, pi. 5, fig. 13)

SYNONYM: Chaetoceros septentrionalis auct. non 0strup (1895), sensu Cleve (1896: 9, pi. 2, fig. 8)

ADDITIONAL references: Hustedt (1927-30), Cupp (1943: fig. 101), Priddle & Fryxell (1985:
A-F)
48, figs

Fig. 2.14f

description: Cells solitary, rectangular in girdle view, isovalvate; pervalvar axis 6-10 urn.
Valves elliptical; apical axis 6-10 um. Setae fine, smooth, diverging at 60-90° to the
pervalvar axis. Chloroplasts 2 per cell. Resting spores discoid.

distribution: Weddell-Scotia Sea (Buck & Garrison, 1983; Garrison et al, 1987); Ross Sea
(Watanabe, 1982); Southern Ocean (Priddle & Fryxell, 1985); Bransfield Strait (Priddle,
1985).

Chaetoceros neogracilis has also been reported from the Mediterranean, Baltic and Arctic seas.

Chaetoceros pendulus G.Karst. (Karsten, 1905: 118, pi. 15, fig. 7)

ADDITIONAL references: Hendey (1937: 295), Priddle & Fryxell (1985: 38, figs A-G)
Fig. 2.14g

description: Cells solitary, heterovalvate; pervalvar axis 8-12 pm. Upper valve with a
central depression; lower valve with projecting corners; apical axis 9-18 um (18-28 \im,fide
Manguin, 1960). Setae long, smooth, orientated downwards in a long sweeping curve.
Chloroplasts small, numerous, extending well into the setae.

distribution: Weddell-Scotia Sea (Garrison et al., 1987); Ross Sea (Watanabe, 1982);
Southern Ocean (Priddle & Fryxell, 1985); Bransfield Strait (Priddle, 1985); South Shetlands
to Cape Horn (Hendey, 1937).

Hasle & Syvertsen (1997: 193) noted that C. pendulus is possibly conspecific with
C. aequatorialis Although there are many similarities between the two, C. aequatorialis has
.

robust, spinose setae, an apical axis of c. 25 pm, and it occurs in temperate and warm waters.
By contrast, C. pendulus, with smooth setae, is a cold-water species known from several
records in Antarctic and sub-Antarctic waters.

Chaetoceros peruvianus Brightw. (Brightwell, 1856: 107, pi. 7, figs 16-18)

SYNONYM: Chaetoceros chilensis Krasske (1941: 266, pi. 4, fig. 3)

See VanLandingham (1968: 736) for full synonymy.

ADDITIONAL REFERENCES: Manguin (1954: 15), Hustedt (1958: 136), Manguin (1960: 288, pi. 1 1,
figs 116, 117), Koch & Rivera (1984: 69, figs 36^17), Priddle & Fryxell (1985: 40), Hasle &
Syvertsen (1997: 195, pi. 38, tab. 46)

Fig. 2.13b

36
 DIATOMS

description: Cells usually solitary, sometimes in short chains, rectangular in girdle view,
with a pronounced constriction between valve mantles and the narrow girdle, heterovalvate;
pervalvar axis 13-20 urn; apical axis 10-32 urn (10-44 urn, fide Hendey, 1937). Upper valve
rounded, with setae arising as a vault-like structure from the centre; lower valve flat with
setae arising from inside the valve margin; small central spine present. All setae curving
back towards lower end of chain and strongly spinose. Chloroplasts small, spherical.

DISTRIBUTION: South America, South Africa, Falkland Is., Humboldt Current (Manguin,
1960); Weddell Sea (Garrison & Buck, 1989b); Southern Ocean (Manguin, 1954; Priddle &
Fryxell, 1985); Ross Sea (Watanabe, 1982); Bransfield Strait (Priddle, 1985; Kopczyhska &
Ligowski, 1985); Wilkes Land, Antarctica (Bunt & Wood, 1963); South Atlantic between
41°30'S 0°01'W and 39°14'S 0°03'W; Antarctic, 66°32'S 0°14'W (Hustedt, 1958).

Chaetoceros radicans Schutt (1895: 48, fig. 27)

ADDITIONAL REFERENCES: VanLandingham (1968: 739), Priddle & Fryxell (1985: 50, figs A-H),
Hasle & Syvertsen (1997: 213, pi. 45, tab. 52)

Fig. 2.13c

DESCRIPTION: Cells small, in straight or curved chains, twisted about the chain axis,
isovalvate, rectangular in girdle view; pervalvar axis 8-15 urn. Valves elliptical; apical axis
6-25 um. Apertures between cells narrow, elliptical, with a central constriction. Setae long,
from narrow ends of valve margins, diverging and lying perpendicular to the cell
thin, arising
chain; intercalary setae strongly spinose; terminal setae smooth. Resting spores in pairs, the
parent cells with fused setae and no aperture.

distribution: Southern Ocean, south of Australia; Southern Ocean (Priddle & Fryxell,
1985); cosmopolitan (Hasle & Syvertsen, 1997).

Chaetoceros simplex Ostenf. (Ostenfeld, 1901: 137, fig. 8)

additional references: VanLandingham (1968: 743), Hendey (1964: 137, pi. 19, fig. 2),
Hasle & Syvertsen (1997: 223, pi. 47, tab. 55)

Fig. 2.13d

description: Cells small, solitary, rectangular in girdle view, isovalvate; pervalvar axis c.
10 um. Valves elliptical; apical axis 4-30 urn. Setae long, thin, arising from narrow ends of
valve surface and lying in the direction of the apical axis. Chloroplasts small, numerous.

distribution: Coastal waters off Davis Station, East Antarctica; Antarctica (Bunt & Wood,
1963); Weddell Sea (Buck & Garrison, 1983); Ross Sea (Watanabe, 1982).

Chaetoceros socialis Lauder (1864: 77, pi. 8, fig. 1)

ADDITIONAL REFERENCES: Cupp (1943: 143, fig. 100), VanLandingham (1968: 744), Priddle &
Fryxell (1985: 52, figs A-I), Hasle & Syvertsen (1997: 221, pi. 47)

Fig. 2.14h

description: Cells in short curved chains, sometimes in spherical colonies, isovalvate.

Valves elliptical; apical axis 2-14 um. Apertures between cells slit-like, the corners of
adjacent valves not touching those of the adjacent cell. Setae fine, each cell having 3 short
and 1 long setae, the long setae of adjacent cells entwining. Chloroplast single.

distribution: Bransfield Strait (Kopczyhska & Ligowski, 1985); Ross Sea (Watanabe,
1982); Southern Ocean (Priddle & Fryxell, 1985).

Chaetoceros tortissimus Gran (1900: 122, pi. 9, fig. 25)

ADDITIONAL references: Cupp (1943: 142, fig. 99), VanLandingham (1968: 749), Hasle &
Syvertsen (1997: 215, pi. 45, tab. 52)

37
 DIATOMS

Fig. 2.14i

description: Cells small, in straight or curved chains, strongly twisted about the chain axis,
rectangular in girdle view; pervalvar axis c. 17 um. Valves elliptical; apical axis 11-20 urn.
Apertures between cells usually only visible at cell corners. Setae long, thin, arising inside of
valve margin, orientated in all directions. Resting spores unknown.

distribution: Southern Ocean (Priddle & Fryxell, 1985); Liitzow-Holm Bay (Tanimura et
al, 1990); Ross Sea (Watanabe, 1982); Bransfield Strait (Kopczyhska & Ligowski, 1985);
cosmopolitan (Hasle & Syvertsen, 1997).

Family Coscinodiscaceae

Coscinodiscus asteromphalus Ehrenb. (Ehrenberg, 1844a: 77)

SYNONYMS: Coscinodiscus radiatus var. asteromphalus (Ehrenb.) Ehrenb. (Ehrenberg, 1854: pi. 18, fig. 45)
Coscinodiscus asteromphalus var. conspicua Grunow, in van Heurck (1883: 130-131, figs 2-5)
Coscinodiscus asteromphalus var. genuina Grunow (1884: 78), nom. inval.

Coscinodiscus oculus-iridis auct. non Ehrenberg (1854), sensu Grunow (1898), in Schmidt et al. (1874-1959:
pi. 63, fig. 5)

ADDITIONAL REFERENCES: Hustedt (1927-30: 452, fig. 250), Hendey (1937: 243), Fryxell &
Hasle (1973: 79, figs 38, 39), Ferreyra & Ferrario (1983b: 1, pi. 1-4), Priddle & Fryxell (1985:
128, figs A, B), Hasle & Lange (1992: 42, figs 1-14), Hasle & Syvertsen (1997: 104, pi. 15)

Fig. 2.15a-d

description: Cells solitary, discoid; pervalvar axis 20-35 pm. Valves 80-400 um diameter,
slightly convex, with a small central depression; mantles gently sloping. Central rosette of
areolae present; areolae arranged in radial striae, polygonal, with cribra, 3-5 in 10 um in the
centre, 6 in 10 um at the margin. Central labiate processes absent; row of marginal labiate
processes 1-3 in 10 um; 2 large curved labiate processes nearer margin (Fig. 2.1 5d); second
row of labiate processes on mantle. Chloroplasts large, numerous, plate-like.
distribution: Coastal waters off Davis Station, East Antarctica; Terre Adelie (Manguin,
1960); Loubet Coast, Antarctic Peninsula (Ferreyra & Ferrario, 1983b); South Atlantic,
Antarctic sector, near 66°32'S 0°14'W (Hustedt, 1958); South Orkney Is. (Ferreyra &
Ferrario, 1983b); coastal waters off Syowa Station, East Antarctica (Ishikawa et al, 2001).

Coscinodiscus bouvet G.Karst. (Karsten, 1905: 83, pi. 3, fig. 9a, b)

ADDITIONAL REFERENCES: Manguin (1915: 52, fig. 36), Heiden & Kolbe (1928: 495), Hendey
(1937: 244-245, pi. 13, figs 3, 4), Priddle & Fryxell (1985: 130, figs A, B), Everitt & Thomas
(1986: 5, fig. 3), Priddle & Thomas (1989: 163-165, figs 1-10), Hasle & Syvertsen (1997: 109,
pi. 17, tab. 20a, b)

Fig. 2.16a-f

description: Cells solitary or in chains of 2-18 cells, characteristically octagonal in girdle

view; pervalvar axis 144-180 um (Fig. 2.16d); girdle broad. Valves circular, 140-200 um
diameter, flat; central part raised, c. 50-75% of total diameter. Central rosette of areolae
distinct (Fig. 2.16a, b); areolae hexagonal, 6 in 10 um, decreasing slightly in size toward
margin. Central labiate processes absent; row of marginal labiate processes 2-3 in 10 um.
Chloroplasts numerous, irregular, stellate.

DISTRIBUTION: Wide distribution in Antarctic waters and southern seas; Liitzow-Holm Bay
(Tanimura et al., 1990); South Georgia, the Sandwich Group, Bransfield Strait, Cape of
Good Hope (Hendey, 1937); inshore Davis Station, Prydz Bay, East Antarctica (Everitt &
Thomas, 1986).

Coscinodiscus centralis Ehrenb. (Ehrenberg, 1844a: 78)

SYNONYMS: Coscinodiscus asteromphalus var. centralis (Ehrenb.) Grunow (1884: 79)

38
 DIATOMS

Figure 2.14. Chaetoceros species, a, C. brevis, Chain of cells in broad girdle view, b, C.
concavicornis, Chain of cells in broad girdle view, c, C. curvisetus, Chain of cells in broad
and narrow girdle views, d, C. debilis, Typical chain of cells, e, C. laciniosus, Typical
colony of cells, f, C. neogracilis, Cell in broad girdle view and apical view, g, C. pendulus,
Solitary cells in broad girdle view, h, C. socialis, Typical colony of cells, i, C. tortissimus,
Chain of cells mostly in broad girdle view. Scale bars: a-i = 10 um. a-i after Cupp (1943).

39
 DIATOMS

Figure 2.15. Coscinodiscus asteromphalus. a, SEM Solitary cell in valve view, b, SEM Cell
in oblique pervalvar view. c. SEM Polygonal areolae with overlaying cribra. d, SEM Internal
valve view with 4 small marginal labiate processes, and large labiate process. Scale bars: a,
1

b = 50 urn; c, d = 10 um.

40
 DIATOMS

Figure 2.16. Coscinodiscus bouvet. a, LM Valve view, focussed on top of raised area of
valve, b, SEM Girdle view, c, SEM
Valve view, focussed on sloping edges of valve, d, SEM
Valve view internal; note marginal labiate processes, e, SEM Areolae forming the central
rosette, f, SEM Valve view internal, enlarged section of d, detail of cribra. Scale bars: a-d =
10 (xm; e, f = 5 um

41
 DIATOMS

Figure 2.17. Coscinodiscus oculoides. a, SEM Valve view. b. SEM Marginal ring of small
labiate processes (arrows), c, SEM Detail of external areolar surface, d, SEM Internal view
of areolae, 2 small labiate processes (arrows) and 1 larger labiate process midway between
smaller processes. Scale bars: a, b = 10 urn; c, d = 1 urn.

42
 DIATOMS

Coscinodiscus oculus-iridis var. tenuistriata Grunow (1884: 77)

See VanLandingham (1968: 873) for full synonymy.

ADDITIONAL REFERENCES: Ehrenberg (1854: pi. 18, fig. 39; pi. 21, fig. 3; pi. 22, fig. 1), Hendey
(1964: 245), Priddle & Fryxell (1985: 132, figs A, B), Hasle & Syvertsen (1997: 105, fig. 6a-c;
pi. 15, tab. 20a, b)

description: Cells solitary, discoid. Valves circular, slightly convex, 100-300 urn diameter;
pervalvar axis 50-60 [im (Hendey, 1964). Central rosette of large areolae conspicuous.
Surrounding areolae 4 6 in 10 urn, becoming finer (6 in 10 urn) towards the margin, arranged
radially in spiralling rows. Marginal ring of labiate processes within 3 or 4 areolae the margin,
visible by LM. Most processes small, but 2 are comparatively large; processes 2-5 areolae
apart. Chloroplasts small, plate-like.

distribution: Southern Ocean (Priddle & Fryxell, 1985); South Georgia (Hendey, 1964);
Bransfield Strait (Priddle, 1985); Ross Sea (Watanabe, 1982).

Coscinodiscus curvatulus Grunow (1878), in Schmidt et al. (1874-1959, pi. 57,

figs 33-35)

See VanLandingham (1968: 883) for full synonymy.

additional references: Mann (1937: 40), Priddle & Fryxell (1985: 150, figs A, B)

Fig. 2.18a

description: Cells (presumably) discoid. Valves circular, 40-100 \im diameter. Areolation
fasciculate; areolae 6-10 in 10 urn, irregular in central area, otherwise in curved rows.
Marginal band of small areolae narrow, striate; striae 12-16 in 10 urn (Hustedt, 1927-30).
Single ring of marginal labiate processes; one process at the end of leading row of fascicle.

distribution: Antarctic, near Commonwealth Bay, East Antarctica (Mann, 1937); South
Atlantic and Scotia Seas (Hendey, 1937); Southern Ocean (Priddle & Fryxell, 1985).
This species has rarely been reported from the Antarctic. Priddle & Fryxell (1985) noted
that
its generic position is uncertain, and that it may belong to Actinocyclus (Hemidiscaceae).

Coscinodiscus horridus G.Karst. (Karsten, 1905: 78, pi. 5, figs 9, 9a)

description: Cells (presumably) discoid. Valves circular, 82-86 um diameter. Areolation

fine, radially fasciculate; marginal band of areolae 4-5 um wide. Spines in 2 regular rows
near the marginal band; spines of outer row longer than inner.

distribution: Southern Ocean, Indian sector (Karsten, 1905).

Priddle & Fryxell (1985) reported and illustrated C. horridus from the Southern Ocean, but
they noted that the material was probably referrable to Thalassiosira (Thalassiosiraceae).

Coscinodiscus marginatus Ehrenb. (Ehrenberg, 1844a: 78)

See VanLandingham (1968: 923) for full synonymy.

ADDITIONAL REFERENCES: Ehrenberg (1854: 18, fig. 44), Hendey (1964: 78, pi. 22, fig. 2), Hasle
& Syvertsen (1997: 107, pi. 18, tab. 20a, b)

Fig. 2.18b

description: Cells solitary, discoid, thick-walled. Valves circular, flat, 35-200 |im diameter.
Central rosette of areolae absent. Areolae coarse, 1-4 in 10 um, in irregular radial rows.
Marginal area prominent, with a ring of labiate processes (not visible by LM) 2-10 areolae
apart. Chloroplasts small, spherical.

distribution: Ross Sea (Watanabe, 1982); Bransfield Strait (Boden, 1985); Amundsen Sea,
70°50'S 102°13'W, "Belgica" (van Heurck, 1909).

43
 DIATOMS

Coscinodiscus oculoides G.Karst. (Karsten, 1905: 81, pi. 6, fig. 3)

ADDITIONAL REFERENCES: van Heurck (1909: 49, pi. 12, fig. 167), Hustedt (1958: 116, figs 32,
33), Kopczyriska & Ligowski (1985), Priddle & Fryxell (1985: 136, figs A, B), Moisan & Fryxell
(1993:493)

Fig. 2.17a-d

description: Cells solitary, discoid. Valves circular, strongly convex, sometimes flattened
towards the centre, 87-230 urn diameter. Central rosette of areolae absent. Areolae
hexagonal with a small central puncta, uniform in size on valve surface, 3-5 in 10 urn.
Marginal labiate processes small, 1 in 10 urn, in a marginal ring (Fig. 2.17c); one (?or 2)
larger marginal process present (Fig. 2. 1 7d). Chloroplasts small, spherical or discoid.

distribution: Coastal waters near Davis Station, East Antarctica; widespread throughout the
Southern Ocean (Hendey, 1937); Prydz Bay, East Antarctica (Ligowski, 1983); Ross Sea
(Watanabe, 1982); Bellingshausen Sea, 69°21'S 81°08'W, "Belgica" (van Heurck, 1909).

Hendey (1964) noted a resemblance to C. oculus-iridis. However, the latter has a

conspicuous, central rosette of c. 5 areolae.

Coscinodiscus oculus-iridis (Ehrenb.) Ehrenb. (Ehrenberg, 1854: pi. 18, fig. 42; pi. 19,

fig- 2)
BASIONYM: Coscinodiscus radiatus var. oculus-iridis Ehrenb. (Ehrenberg, 1839: 147), Ehrenberg (1854: pi. 18,
fig. 42; pi. 19, fig. 2), van Heurck (1896: 530)

SYNONYMS: Coscinodiscus oculus-iridis var. genuina Grunow (1884: 77), nom. inval.
Coscinodiscus oculus-iridis var. typicus Cleve-Euler (1942: 258, fig. 30), nom. inval.

See VanLandingham (1968: 937) for full synonymy.

ADDITIONAL REFERENCES: Hustedt (1927-30: 454, fig. 252), Hendey (1937: 249; 1964: 78, pi. 24,
fig. 1), Hargraves (1968: 138, fig. 139A, B), Priddle & Fryxell (1985: 138, figs A, B)

Fig. 2.18c

description: Cells solitary, discoid. Valves circular, flat or slightly convex, 180-260 urn
diameter. Central rosette of c. 5 areolae, conspicuous. Areolae small near the centre,
increasing in size towards the margin, arranged radially. Chloroplasts small, spherical.

distribution: South Georgia (Hendey, 1937); Ross Sea (Watanabe, 1982); Weddell Sea
(Garrison & Buck, 1989b); Bransfield Strait (Priddle, 1985); Weddell-Scotia Sea (Garrison
etal., 1987).

Coscinodiscus oppositus G.Karst. (Karsten, 1905: 82, pi. 7, fig. 5)

ADDITIONAL REFERENCE: Priddle & Fryxell (1985: 154, fig. on p. 55)

description: Cells (presumably) discoid. Valves circular; apical axis 44-90 urn, with
conspicuous marginal processes and, apparently, 2 radial labiate processes. Valve margins
with small spines.

distribution: Southern Ocean (Hendey, 1937; Priddle & Fryxell, 1985).

This species has rarely been reported from the Antarctic. Priddle & Fryxell (1985) noted that
its generic position is uncertain.

Coscinodiscus radiatus Ehrenb. (Ehrenberg, 1841: 148, pi. 3, fig. la-c)

See VanLandingham (1968: 950) for full synonymy.

ADDITIONAL REFERENCES: Watanabe (1982), Boden (1985), Hasle & Syvertsen (1997: 107,
fig. 6d, e;pl. 18)

Fig. 2.18d

44
 DIATOMS

Figure 2.18. Coscinodiscus, Actinoptychus, Actinocyclus and Leptocylindrus species, a,

C. curvatulus . b, marginatus. c, C. oculus-iridis. d, C. radiatus. e, C. subtilis. f,
C.
Actinoptychus senarius. g, Actinocyclus adeliae. h, Actinocyclus divisus. i, Z.. danicus. Scale
bars: a-i = 10 urn. a-f after Cupp (1943); g after Manguin (1960); h, i after Hustedt (1958).

45
 DIATOMS

Figure 2.19. Stellar ima microtrias. a, LM Vegetative cell, valve external view, b, SEM
Valve external view, 6 central labiate processes, c, SEM Valve external view, 4 central
labiate processes, d, SEM Same valve as b, external view, e, SEM Valve internal view, 3
central labiate processes, f, SEM Valve internal view (internal lines are artifacts of SEM).
Scale bars: a, c, d, f = 10 urn; b, e = 5 urn.

46
 DIATOMS

description: Cells solitary, discoid. Valves flat, circular, 30-180 urn diameter. Areolation

radial,sometimes indistinctly decussate; areolae 2-9 in 10 urn. Small labiate processes at the
points of origin of incomplete striae. Hyaline lines absent. One marginal ring of processes,
including 2 slightly larger processes c. 135° apart, these visible by LM as indentations of the
valve margin. Smaller processes scarcely visible.

distribution: Ross Sea (Watanabe, 1982); Prydz Bay, SIBEX (Boden, 1985); oceanic
species with world-wide distribution in temperate seas (Hendey, 1937).

Coscinodiscus radiatus has only occasionally been reported from the Antarctic.

Coscinodiscus subtilis Ehrenb. (Ehrenberg, 1841: 412, pi. 1, fig. 3)

See VanLandingham (1968: 971) for full synonymy.

ADDITIONAL references: Karsten (1905: 86, pi. 7, figs 11, 11a), Wood (1960: 218, pi. 3, fig.

15), Bunt & Wood (1963: 1255)

Fig. 2.18e

description: Cells solitary, discoid. Valves flat or convex, 80-130 urn diameter, with fine
hexagonal areolae in fascicles; margin with numerous spinules. Areolation in two systems: a
series of rather more distinct oblique lines crossing radiating fascicles of parallel lines, and
lines parallel to the middle radius of each fascicle (Hendey, 1937). Central area and central
rosette absent. Chloroplasts numerous, plate-like.

distribution: Wide distribution in subpolar seas (Hendey, 1937); Ross Sea (Watanabe,
1982); Antarctic, south of Australia (Wood, 1960; Bunt & Wood, 1963).

Stellarima microtrias (Ehrenb.) Hasle & P.A.Sims (1986: 111, figs 18-27)

BASIONYM: Symbolophora microtrias Ehrenb. (Ehrenberg, 1844b: 205)

SYNONYMS: Coscinodiscus symbolophorus Grunow (1884: 82, pi. 4, figs 3-6)

Coscinodiscus adumbratus 0strup (1895: 461, pi. 8, fig. 90)

Coscinodiscus furcatus G.Karst. (Karsten, 1905: 82, pi. 4, fig. 7), Manguin (1960: 247, pi. 24, figs 288-290),
Priddle & Fryxell (1985: 134, figs A-C)
Coscinodiscus pentas (Ehrenb.) A.Mann (1907: 256)
Coscinosira stellaris (Roper) Heiden var. symbolophora (Grunow) Heiden, in Heiden & Kolbe (1928: 471)
Coscinodiscus (symbolophorus var.?) signatus A.Mann (1937: 46, pi. 4, fig. 8)

Podosira liotardii Manguin (1960: 241, pi. 21, figs 252-254)

Symbolophora furcata (G.Karst.) V.A.Nikolajev (1983: 1 126, pi. 1, figs 8, 9)

ADDITIONAL REFERENCES: Syvertsen (1985: 113-115, figs 1-14), Fryxell (1989: 11); Hasle &
Syvertsen (1990b: 109, pi. 13.1, figs 1, 2), Moisan & Fryxell (1993: 293), Hasle & Syvertsen
(1997: 113, pi. 19)

Fig. 2.19a-f

description: Vegetative cells solitary or chain-forming (see Fryxell, 1989), discoid to

lenticular, attached to sibling cells at the centre or off-centre. Valves flat to slightly convex,
circular, 35-105 urn diameter; centre of valve hyaline, with a ring of up to 8 labiate processes;
processes slit-like externally (Fig. 2.19d), but internally raised and elongated (Fig. 2.19e, f);
marginal processes absent. Areolae arranged in furcate radial striae, 11-16 in 10 um, loculate,
externally simple or with cribra, internally with foramina. Chloroplasts numerous, irregular.
Resting spores 40-100 urn diameter, with 3-5 central labiate processes; areolae fasciculate, 9-
11 in 10 um. See Syvertsen (1985) for details of resting spore formation.

distribution: Type locality, Antarctic sea-ice, 78°10'S 162°W; Haakon VII Sea, 69°05'S
04°25'W and 69°43'S 06°30'E (Hustedt, 1958); Weddell Sea (Fryxell, 1989); near Syowa
Station, East Antarctica (Watanabe et ai, 1990; Ishikawa et ai, 2001); southern cold waters
(Hasle & Syvertsen, 1997); Scotia Sea (Garrison et a/., 1987).

47
 DIATOMS

Stellarima differs from Coscinodiscus in lacking marginal labiate processes, and in having
internal closing plates to the areolae (Round et al., 1990). Resting spores are common in ice
samples and in net samples from the vicinity of ice (Hasle & Syvertsen, 1990b). Hasle &
Syvertsen (1997) observed that "Stellarima mictrotrias forms heavily silicified endogenous
resting spores, usually identified as Coscinodiscus symbolophorus".

Family Eupodiscaceae

Odontella aurita (Lyngb.) C.Agardh (1832: 56)

BASIONYM: Diatoma aurita Lyngb. (Lyngbye, 1819: 182, pi. 62, fig. D)
SYNONYM: Biddulphia aurita (Lyngb.) Breb. & Godey (Brebisson & Godey, 1838: 12)

additional REFERENCES: Cupp (1943: 161, fig. 112), Hasle & Syvertsen (1997: 239, pi. 49, tab.
62)

description: Cells coarsely silicified, solitary or forming short zig-zag chains, rectangular in
girdle view with slightly raised divergent apical corners; pervalvar axis 31-50 um. Valves
elliptical in valve view; valve face between corners evenly inflated; apical axis 10-97 um.
Valve areolae 8-11 in 10 um; girdle band areolae 8-14 in 10 um. Labiate processes 2 per
valve, with long curved external tubes. Chloroplasts numerous, stellate.

distribution: Ross Sea (Watanabe, 1982); Wilkes Land, Antarctica (Bunt & Wood, 1963);
cosmopolitan? (Hasle & Syvertsen, 1997); Prydz Bay, SIBEX (Boden, 1985).

Odontella litigiosa (van Heurck) Hoban, in Hoban et al. (1980: 598, figs 27-38)

BASIONYM: Biddulphia litigiosa van Heurck (1909: 40, pi. 10, fig. 141)
SYNONYM: Biddulphia polymorpha Manguin (1915: 23-27, figs 2-7)
See Hendey (1937: 277) and Manguin (1915: 23-27) for further details of synonymy.

ADDITIONAL REFERENCES: Peragallo (1921: 73), Priddle & Fryxell (1985: 68, figs A-F), Everitt &
Thomas (1986: 8, fig. 3 A), Andreoli et al. (1995: 472, fig. 14)
Fig. 2.20a-f

description: Vegetative cells chain-forming, rectangular in girdle view, with raised corners;
pervalvar axis 60-130 um. Valves oval to elliptical in valve view; apical axis 30-140 um,
with pronounced slightly diverging bipolar ocellate elevations; central area of valve convex.
Valve surface with 2 labiate processes near the centre, poroid, with granules over surface.
Resting spores with weakly or strongly developed hollow spines over the valve surface.
Twinned pores on valve surfaces [described and illustrated in Hoban et al. (1980: figs 28,
35)] are unique to the three phases of the vegetative life cycle (see below).

distribution: Inshore waters near Davis Station, Prydz Bay, East Antarctica (Everitt &
Thomas, 1986); Ross Sea (Andreoliet al, 1995); South Orkney Is. (Hoban et al., 1980).

A polymorphic diatom with three phenotypes: litigiosa, polymorpha and anthropomorpha.

The polymorpha and litigiosa phases probably represent vegetative cells, the anthropo-
morpha phase the resting spore. Intermediate cell types having been described. In the
transition from polymorpha to litigiosa to anthropomorpha, the elevations and spines
elongate, the cell volume decreases, the girdle is reduced, and the valve mantle becomes
constricted in a manner suggestive of resting cell formation (Hoban et al., 1980).

Odontella weissflogii (Grunow) Grunow (1884: 6, 58)

BASIONYM: Biddulphia weissflogii Grunow, in van Heurck (1882: pi. 100, figs 1, 2), Janisch (1888: pi. 21, figs
10-16, as Biddulphia weissflogii Janisch)
SYNONYMS: Biddulphia weissflogii "var.?", Schmidt (1889), in Schmidt et al. (1874-1959. pi. 141, figs 24-26)

Biddulphia striata G.Karst. (Karsten, 1905: 122, pi. 17, figs 2, 3b)
Zygoceros caballeroi Azpeitia (Azpeitia Moros, 191 1: 224, pi. 2, fig. 4)

See Hoban et al. ( 1980) for a discussion of the confused nomenclatural history of O. weissflogii.

48
 DIATOMS

Figure 2.20. Odontella litigiosa. a, LM anthropomorpha phase, valve view, b, SEM

anthropomorpha phase, c,SEM litigiosa phase, oblique valve view with 2 conspicuous
labiate processes, d, SEM litigiosa phase, detail of ocellate elevation, e, SEM
anthropomorpha phase; ocellate elevation with conspicuous granules and hollow spines, f,

SEM anthropomorpha phase, valve internal view. Scale bars: a-f = 10 pm.

49
 DIATOMS

Figure 2.21. Odontella weissflogii. a, LM Chain of vegetative cells, b. SEM Vegetative cell,
girdle view, c, SEMVegetative cell, oblique view of valve, with hollow labiate processes, d,
SEM Vegetative cell, valve internal view of labiate process, e, SEM Resting spore with
reduced ocellations. Scale bars: a, b = 10 urn; c, e = 5 urn; d = 1 urn.

50
 DIATOMS

ADDITIONAL REFERENCES: Hendey (1937: 278, pi. 10, figs 4, 5, as Biddulphia striata), Okuno
(1953: 17), Manguin (1954: 15), Hustedt (1958: 137), Hoban et al. (1980: 594, figs 15-29),
Priddle& Fryxell (1985: 70, figs A-H), Moisan & Fryxell (1993: 293)

Fig. 2.21a-e

description: Cells solitary or forming short chains, weakly silicified, rectangular in girdle
view, with slightly raised corners; pervalvar axis 60-130 um. Valves elliptical in valve view;
apical axis 28-84 um; transapical axis 22-42 um, with 2 or 4 long curved labiate processes,
the external sections of which are hollow (Fig. 2.21b), the internal parts small and simple
(Fig. 2.21c); valves loculate with internal foramina and external cribra; bipolar ocellate
elevations small, mammiform. Chloroplasts numerous, stellate. Resting spores heavily
silicified; valves with reduced bipolar ocellate elevations and only 2 offset labiate processes
that are not hollow in the external sections (Fig. 2.2 le).

distribution: Weddell Sea (Buck & & Fryxell, 1993); Wilkes Land,
Garrison, 1983; Moisan
Antarctica (Bunt & Wood, 1963); Heard
(Manguin, 1954); South Atlantic (Hustedt, 1958);
I.

South Georgia, Bransfield Strait (Hendey, 1937; Kopczyhska & Ligowski, 1985); Lutzow-
Holm Bay (Tanimura et al, 1990); Scotia Sea (Garrison et al., 1987); Kerguelen Basin
(Mangin, 1915).

Family Heliopeltaceae

Actinoptychus senarius (Ehrenb.) Ehrenb. (Ehrenberg, 1843: 400, pi. 1.1, fig. 27;
pi. 1.3, fig. 21)

BASIONYM: Actinocyclus senarius Ehrenb. (Ehrenberg, 1838: 172, pi. 21, fig. 6)

SYNONYMS: Actinocyclus undulatus Kiitz. (Kiitzing, 1844: 132)

Actinoptychus undulatus Bailey ex Ralfs, in Pritchard (1861 : 839, pi. 5, fig. 88)
Actinoptychus undulatus Bailey ex Hustedt (1927-30: 475), nom. illeg.

ADDITIONAL REFERENCES: Bailey (1842: 94, fig. 11, as Actinoptychus sp.), Hendey (1937: 271),
Priddle & Fryxell (1985: 110, figs A-C), Round et al. (1990: 200, fig. A), Hasle & Syvertsen
(1997: 141, pi. 22)

Fig. 2.18f

description: Cells solitary, discoid. Valves divided into 6 sectors, alternately raised and
depressed, 20-150 um in diameter; central area plain and ±hexagonal; remainder of valve with
coarse areolation; areolae 4-7 in 10 um. Each of the raised sectors has a marginal labiate
process with an external tube. Choroplasts numerous, large (Hasle & Syvertsen, 1997).

distribution: Southern Ocean (Priddle & Fryxell, 1985); South Atlantic, Southern Ocean
off Enderby Land, South Georgia, Drake Passage (Hendey, 1937).

Primary valves were formerly described as the genus Debya Pant. (Round et al., 1990).
Hendey (1937) noted that Actinoptychus senarius was often attributed as '''Actinoptychus
undulatus Bailey", but that there were no grounds for this, Bailey's original figure having
been unnamed.

Family Hemidiscaceae

Actinocyclus

Species of Actinocyclus have the following diagnostic features: radial, usually fasciculate
expanded internally; a marginal
areolation; a marginal ring of labiate processes, laterally
pseudonodulus (sometimes difficult to distinguish); and radial, hyaline bands (see Villareal
& Fryxell, 1983 and Watkins & Fryxell, 1986 for further details).

51
 DIATOMS

Actinocyclus actinochilus (Ehrenb.) Simonsen (1982: 105, figs 1-9)

BASIONYM: Coscinodiscus actinochilus Ehrenb. (Ehrenberg, 1844b: 200; 1854: pi. 35A, figs xxi, 5)

SYNONYM: Charcotia actinochilus (Ehrenb.) Hust. (Hustedt, 1958: 126, figs 57-80)

See Simonsen (1982: 105-106) for full synonymy.

ADDITIONAL REFERENCES: Villareal & Fryxell (1983: 461, figs 21-32), Priddle & Fryxell (1985:
102, figs A-C), Everitt & Thomas (1986: 5, fig. 2B), Moisan & Fryxell (1993: 493), Fryxell
(1990: 112, tab. 14.1, fig. 1)

Fig. 2.22a-f

description: Cells solitary, cylindrical to discoid. Valves flat with rounded edges, 20-1 12 urn
diameter. Areolae in isolated radial rows that are sometimes curved or incomplete; areolation
variable; areolae 5-11 in 10 urn, loculate, with a foramen on the inside, bordered by a
thickened rim (Fig. 2.22e, f). Marginal labiate processes 1 in 10 urn, 9-15 urn apart, laterally

expanded (Fig. 2.22e, f). Pseudonodulus sometimes present near junction of flat part of valve
face and mantle, not perforate. Striae on valve mantle 13-21 in 10 um. Chloroplasts
numerous, discoid.

distribution: Coastal waters near Davis Station, East Antarctica (Everitt & Thomas, 1986);
Weddell Sea (Moisan & Fryxell, 1993); Weddell-Scotia Sea (Garrison et al, 1987); Indian
Ocean sector of the Antarctic (Kopczyhska et al., 1986); Kerguelen and Atlantic sectors of
the Antarctic, very common from 68°49'S 0°40'W to 69°43'S 06°30'E, and 69°43'S
06°30'E to 61°32'S 0°00' (Hustedt, 1958); coastal waters off Syowa Station, East Antarctica
(Ishikawae/a/.,2001).

Specimens recorded from near Davis Station have a diameter range of 24-52 urn.

Actinocyclus adeliae Manguin (1957: 1 16, pi. 2, fig. 13a, b)

ADDITIONAL REFERENCE: Manguin (1960: 258, pi. 3, figs 45, 46; pi. 26, figs 308-310)

Fig. 2.18g

description: Cells solitary, discoid. Valves convex in the centre and with rounded edges,
41-68 um diameter. Areolae in c. 13-16 radial (not fasciculate) striae, 12-15 in 10 um.
distribution: Type locality, Terre Adelie, 66°50'S 141°25'E (Manguin, 1957).

Actinocyclus curvatulus Janisch (1878), in Schmidt et al. (1874-1959: pi. 57, fig. 31)

SYNONYMS: Coscinodiscus curvatulus var. subocellatus Grunow (1884: 83, pi. D, fig. 15)

Actinocyclus subocellatus (Grunow) Rattray (1890a: 488; 1890b: 145)

ADDITIONAL REFERENCES: Simonsen (1974: 20), Priddle & Fryxell (1985: 106, figs A, B), Hasle
& Syvertsen(1997: 121, pi. 19)

Fig. 2.23a-d

DESCRIPTION: Cells solitary, discoid, 13-160 um diameter. Areolae fasciculate, in slightly but
distinctly curved rows; areolae 8-18 in 10 um, decreasing
in size towards the margin, with a
distict marginal band formed of small areolae. Marginal processes 12-18 in 10 um, each
located at the end of a row of areolae; processes expanded on internal valve face
(Fig. 2.23d). Pseudonodulus near the margin (Fig. 2.23a), non-perforate.

distribution: Coastal waters near Davis Station, East Antarctica; type locality, Zemlya
Frantsa-Iosifa (Franz Josef Land), Arctic Ocean (Grunow, 1884); Kerguelen Is. (Grunow,
o
1884); South Atlantic, 46°32'S 0°02'W-49 00'S 0°07'W (Hustedt, 1958).

The rows are an especially distinctive feature of this

fasciculate areolae in slightly curved
species. Valve diameter of A. curvatulus seems to be rather variable, e.g. 36-57 um (Hustedt,
1958) and 124-160 um (Mann, 1937). Cells collected near Davis Station were 25-40 um in
diameter, with fasciculate areolae 8 in 10 um in the centre, c. 15 in 10 um and decreasing in
diameter towards the margin, and labiate processes 1-2 in 10 um in a marginal ring.

52
 DIATOMS

Figure 2.22. Actinocyclus actinochilus. a, LM Valve face with areolae, b, SEM Valve face
with 8 marginal labiate processess and indistinct areolae, c, SEM Valve face with areolae, d,
SEM Oblique view of cell showing fine striation of valve mantle and marginal
pseudonodulus. e, f, SEM Inner surface of valve showing laterally expanded internal ends of
the labiate processes. Scale bars: a-f = 5 pm.

53
 DIATOMS

Figure 2.23. Actinocyclus curvatulus. a, SEM Valve face with 10 marginal labiate
processess and pseudonodulus (arrow), b, SEM Valve face with partially etched areolae, c,
SEM Valve face with fasciculate areolae and 12 marginal labiate processes, d, SEM Internal
valve surface showing laterally expanded internal ends of the labiate processes. Scale bars:
a-d = 10 uni.

54
 DIATOMS

Actinocyclus divisus (Grunow) Hust. (Hustedt, 1958: 129, fig. 81)

BASIONYM: Coscinodiscus divisus Grunow (1878: 125; 1884: 83, pi. D, fig. 16)

SYNONYMS: Coscinodiscus {curvatulus var.?) divisus Grunow (1884: 31, pi. 4, fig. 16)

Actinocyclus cholnokyi (Grunow) VanLand. (VanLandingham, 1967: 83)

ADDITIONAL REFERENCES: Hustedt ( 958: 129, figs 82, 83), Priddle

1 & Fryxell (1985: 104, fig. A),
Fryxell(1990: 112, tab. 14.1), Moisan & Fryxell (1993: 493)

Fig. 2.18h

description: Cells solitary, cylindrical. Valves flat, with rounded edges, 41-70 urn diameter.
Striae on valve mantle forming slightly fasciculate decussate rows, 20-24 in 10 urn. Areolae
7-9 in 10 pm. Labiate processes 8-15 urn apart, laterally expanded. Pseudonodulus
marginal, areolate or eroded.

distribution: Antarctic, cosmopolitan (Fryxell, 1990); South Atlantic (Hustedt, 1958).

There is confusion regarding the correct name of this species. Simonsen (1974) considered
A. divisus to be conspecific with A. curvatulus, whereas VanLandingham (1967) regarded it
as a synonym of A. cholnokyi, and Fryxell et al. (1986) included it in the synonymy of
Azpeitia neocrenulata (VanLand.) G. A. Fryxell & Watkins.

Actinocyclus exiguus G. A. Fryxell & Semina (1981: 442, figs 1-10)

ADDITIONAL REFERENCES: Fryxell (1990: 112, pi. 14.1, fig. 2), Hasle & Syvertsen (1997: 122)

Fig. 2.24a-f

description: Cells solitary, cylindrical. Valves circular, flattened, with a high mantle and a
thin delicate fringing skirt that partly overlaps the first girdle band; apical axis 6.5-15 urn.
Areolae rows, sometimes fasciculate, 12-18 in 10 pm in the centre, 21-24
in irregular radial
in 10 umtowards the margin. Labiate processes slightly expanded, 3-5 per valve, located on
mantle, c. 1 um long and laterally expanded (viewed from inside of valve). Pseudonodulus
sometimes visible, possibly areolate.

distribution: Southern Ocean, south of Australia; type locality, Southern Indian Ocean,
46°04.7'S 49°19.0'E (Fryxell & Semina, 1981); Antarctic and sub-Antarctic (Fryxell, 1990).

Actinocyclus octonarius Ehrenb. (Ehrenberg, 1838: 172, pi. 21, fig. 7)

SYNONYM: Actinocyclus ehrenbergii Ralfs, in Pritchard (1861: 834)

See VanLandingham (1967: 100) for further details of synonymy.

ADDITIONAL REFERENCES: Hustedt (1958: 130), Priddle & Fryxell (1985: 108, fig. on p. 109),
Ricard (1987: 43, figs 35-39), Hasle & Syvertsen (1997: 120)

DESCRIPTION: Cells solitary, discoid. Valves slightly convex, to 50-300 um diameter.

Areolation fasciculate and with rows parallel to central row of fascicle, sometimes with
hyaline areas adjacent to radial edge of fascicle. Areolae 6-8 in 10 pm, but much finer at the
margin. Pseudonodulus at the junction of the valve face and mantle, distinct, operculate. A
marginal labiate process is located at the end of each radial row of a fascicle.

distribution: Southern Ocean (Priddle & Fryxell, 1985); cosmopolitan (Hasle & Syvertsen,
1997); South Atlantic, c. 49-59°S 0°00'W (Hustedt, 1958).

Actinocyclus octonarius is the type species of the genus. This has rarely been recorded from
polar waters.

Actinocyclus spiritus Watkins, in Watkins & Fryxell (1986: 310, figs 17-23)

ADDITIONAL REFERENCES: Fryxell (1990: 112, tab. 14.1, pi. 14.1, fig. 3), Hasle & Syvertsen
(1997: 122, tab. 26)

55
 DIATOMS

description: Cells solitary, cylindrical to discoid. Valves with rounded edges, 27-36 um
flat,

diameter. Areolation variable; areolae in curved or incomplete rows, 13-15 in 10 um,

abruptly changing to 24-29 in 10 um at the margin, loculate, with a foramen on the inside,
bordered by a thickened rim. Hyaline margin absent. Marginal labiate processes 8-12,
slightly expanded, 10-12 um apart. Chloroplasts numerous, discoid.

distribution: Southern Ocean, 62°05.4'S 39°05.6'W, AMERIEZ 83, Station 16 (Watkins &
Fryxell, 1986); Antarctic ice-edge (Fryxell, 1990); Southern Ocean, Indian sector (Waters et
al, 2000).

The rare Actinocyclus spiritus is morphologically most similar to A. actinochilus, the latter
having coarser valve areolation (5-1 1 in 10 um).

Azpeitia nodulifera (A. W.F.Schmidt) G.A. Fryxell & P.A.Sims, in Fryxell et al. (1986:
19, figs 17, 18.1-18.5,30.3,30.4)

BASIONYM: Coscinodiscus nodulifer A. W.F. Schmidt (1878), in Schmidt et al. (1874-1959: 15, pi. 59, figs 22,
23) (fide Rattray, 1890a: 520)
SYNONYMS: Coscinodiscus radiatus H.L.Sm. (Smith, 1874-79: no. 98)

Coscinodiscus nodulifer var. apiculata A. W.F. Schmidt (1878), in Schmidt et al. (1874-1959: 15, pi. 59,
fig. 20) (fide Rattray, 1890a: 520)

ADDITIONAL REFERENCES: DeToni (1894: 1249), Mann (1907: 255), Karsten (1907: 264, pi. 36,

fig. 6), Boyer (1926-27: 55), Hustedt (1927-30: 426, fig. 229), Hendey (1937: 248; 1964: 77, pi.
22, fig. 10), Fryxell & Hasle (1972: 69), Hasle & Syvertsen (1997: 126, pi. 21)

Fig. 2.25f

DESCRIPTION: Cells solitary, cylindrical; pervalvar axis 17-18 um. Valves flat, circular;
apical axis 20-102 um. Areolae in radial rows, 3-8 in 10 um, each areola surrounded by 6
small granules giving a hexagonal pattern to valve; areolae on mantle in 2 rows under a
thickened ridge; areolae adjacent to the ridge tear-drop-shaped, those beside the hyaline
margin elongated pentagons. Striae on mantle 5-8 in 10 um. Central labiate process with a
tubular rim; marginal processes 1-2 in 10 um, internally perpendicular to radius.

distribution: Southern Ocean, south of Australia; Prydz Bay, East Antarctica (Ligowski,
1983); type locality, Gulf of Mexico (Schmidt, 1878); Ross Sea (Watanabe, 1982).

Rattray (1890a) based Coscinodiscus nodulifer var. apiculata on Schmidt's illustration

(1878, pi. 59. fig. 20), of a partial valve but one that clearly shows the irregular
which is

radial areolation, and periodicity around the margin as in the modern

central process,
circumscription of A. nodulifera. Here we include C. nodulifer var. apiculata of Schmidt
(1878, fig. 20) in synonymy with A. nodulifera.

Azpeitia tabularis (Grunow) G.A. Fryxell & P.A.Sims, in Fryxell et al. (1986: 16,
figs 14, 15,30,31)
BASIONYM: Coscinodiscus tabularis Grunow (1884a: 86)
SYNONYM: Coscinodiscus tabularis var. antarctica Manguin (1960: 251, pi. 24, figs 293-295)

ADDITIONAL REFERENCES: Hustedt (1927-30: 427, fig. 230a), Manguin (1954: 19), Hustedt
(1958: 119, figs 48-56), Priddle & Fryxell (1985: 124, figs A, B), Fryxell (1990: 112, pi. 14.1,
fig. 4), Hasle & Syvertsen (1997: 126, pi. 20)

Fig. 2.25a-e

description: Cells solitary, discoid; pervalvar axis c. 7 um. Valves circular, flat; apical axis
16-70 um. Areolae slightly fasciculate, in radial rows, 5-10 in 10 um near the centre, c. 16
in 10 um towards the margin. Labiate processes in a marginal ring, 7-12 in 10 um, laterally
expanded, with a reduced neck (Fig. 2.25b). Labiate process solitary, large, located on the
edge of the central annulus. Pseudonodulus absent.

distribution: Coastal waters near Davis Station, East Antarctica; type locality, Tafelkai,

56
 DIATOMS

Figure 2.24. Actinocyclus exiguus. a, LM Solitary cell, b, SEM Oblique view of cell with
conspicuous labiate processes, c, SEM Valve view of cell with 4 marginal labiate processes.
d, SEM Oblique internal view of valve with expanded internal ends of the labiate processes.
e, SEM Internal view of same cell as d, of valve and girdle band, f, SEM Internal view of

eroded valve. Scale bars: a-f =

1 urn.

57
 DIATOMS

Figure 2.25. Azpeitia species, a-e, A. tabularis. a, LM Vegetative cell showing coarse
areolation. b, SEM Valve, internal view; note laterally expanded marginal and near-central
labiate processes, c, SEM Valve, external view; note marginal and near-central labiate
processes, d, SEM Oblique external view of valve; note marginal labiate processes and valve
mantle, e, SEM Valve, oblique external view of valve showing 3 labiate processes and girdle
bands, f, A. nodulifera, SEM Oblique view of vegetative cell; note central tubular rim of
central process. Scale bars: a-d, f = 10 um; e = 1 urn.

58
 DIATOMS

Zemlya Ocean (Grunow, 1884); Prydz Bay, East

Frantsa-Iosifa (Franz Josef Land), Arctic
Antarctica (Ligowski, 1982); in Euphausia from Kerguelen
1983); Ross Sea (Watanabe,
sector of the Southern Ocean, in Salpa spp. from 68°49'S 0°40'W to 46°32'S 0°02'W,
common around 51°S, in plankton in the South Atlantic, from 51°29'S 0°08'W to 46°32'S
0°02'W (Hustedt, 1958).
Fryxell et al. ( 1986) noted a wide range of morphological variation in this species.

Hemidiscus cuneiformis G.C.Wall. (Wallich, 1860: 42, pi. 2, figs 3, 4)

SYNONYMS: Euodia cuneiformis (G.C.Wall.) Scluitt (1896: 100, fig. 174D, E)

Euodia inornata Castrac. (Castracane, 1886: 149, pi. 12, fig. 1)

Euodia radiata Castrac. (Castracane, 1886: 150, pi. 12, fig. 4)

Figure 2.26. Hemidiscus cuneiformis. a, LM Vegetative cell, b, SEM Valve, external view.
c, SEM Oblique view of ventral margin of valve; note marginal labiate process, d, SEM
Oblique view of inflated dorsal margin of valve, e, SEM Detail of pseudonodulus and
surrounding areolae, f, SEM Valve, internal view of areolae and expanded end of labiate
process. Scale bars: a-d = 10 um; e, f = um. 1

59
 DIATOMS

ADDITIONAL REFERENCES: Hendey (1937: 264; 1964: 94, pi. 22, fig. 9), Fryxell & Hasle (1973:
76, figs 18, 19), Hasle & Syvertsen (1997: 128, pi. 22, tab. 28)

Fig. 2.26a-f

DESCRIPTION: Cells solitary(?), wedge-shaped. Valves flat, almost semicircular; dorsal

margin strongly convex; ventral margin weakly convex, often with a median inflation on the
ventral side; apical axis 40-174 um; transapical axis 30-90 urn. Areolae in irregular radial
fascicles, but in short parallel lines the towards the valve centre where 6-12 in 10 um, and in
irregular or tangential sectors towards the margin and poles where 10-18 in 10 um. Poles
with small inconspicuous processes; central hyaline area and rosette absent; ventral margin
of valve with small spinules and a small partially occluded pseudonodulus midway along
margin. Labiate processes 1-2 in 10 um along ventral margin only, with 1 on either side of
the pseudonodulus. Girdle simple, lacking intercalary bands. Chloroplasts numerous, small,
spherical.

distribution: Coastal waters near Davis Station, East Antarctica; South Atlantic,
47°19.75'S 1 1°05'E to 49°50'S 08°32.5'E (Hendey, 1937).

Hendey (1937) referred all specimens, including a series of intermediate forms which "made
it impossible to recognise the value of nomenclatural species and varieties", to
H. cuneiformis. The genus now includes only two species, H. cuneiformis and the warm-
water H. kanayanus Simonsen (Simonsen, 1972).

Roperia tesselata (Roper) Grunow ex Pelletan (1889: 158, fig. 417)

BASIONYM: Eupodiscus tesselatus Roper (1858: 19, pi. 3, fig. la, b)

SYNONYMS: Actinocyclus coscinodiscoides A.Mann (1937: 19, pi. 1, fig. 5)

Roperia tesselata var. coscinodiscoides (A.Mann) Kolbe (1955: 176)

ADDITIONAL REFERENCES: Simonsen (1974: 22), Fryxell et al. (1986: 24, figs 25, 32.3, 32.4),
Ricard (1987: 43, figs 40^6), Hasle & Syvertsen (1997: 130, pi. 22)

Fig. 2.27a-c

description: Cells discoid. Valves circular to oval; apical axis 25-70 um. Areolation
irregular to eccentric, 6-7 in 10 um at valve centre, smaller and c. 14 in 10 um near the
margin. Marginal labiate processes 1-2 in 10 um. Pseudonodulus prominent, near the margin
and close to one labiate process, operculate. Chloroplasts numerous, small.

distribution: Coastal waters near Davis Station, East Antarctica; sub-Antarctic Pacific, to
57°S (Hasle, 1976a).

The cells found near Davis Station measured 25-40 um in diameter. In all cases, the
pseudonodulus is operculate, thus belonging to the "simplex" type of Ross & Sims (1972),
i.e. "Type C" of Simonsen (1975: 83, text fig. 1).

Family Leptocylindraceae

Corethron inerme G.Karst. (Karsten, 1905: 104, pi. 13, figs 11-17)
SYNONYMS: Corethron criophilum "inerme" phase (G.Karst.) Hendey (1937: 329, pi. viii, fig. 9)

Corethron criophilum f. inerme (G.Karst.) Hasle (1969: 83)

ADDITIONAL REFERENCES: Hustedt (1958: 131), Priddle & Fryxell (1985: 54, figs A, B), Thomas
&Bonham(1990: 105, pi. 12, figs 2, 4, 6), Crawford et al. (1998: 1 1, figs 3, 44-59)

Fig. 2.28a-f

description: Cells chain-forming, weakly silicified, cylindrical; pervalvar axis 40-350 um.
Valves weakly convex, circular; apical axis 20-62 um; only end cells of colonies with
dissimilar valves; intercalary valves with short spines interlocking siblings. Upper valve of

60
 DIATOMS

Figure 2.27a-c. Roperia tesselata. a, SEM Valve face with fasciculate areolae and pseudo-
nodulus. b, SEM Internal valve surface with marginal labiate processes and pseudonodulus.
c, SEM Internal opening of the pseudonodulus. d, e, Guinardia cylindrus. d, SEM Single cell
with attachments to two (broken) cells, e, SEM Detail of central labiate processes attaching
adjacent cells. Scale bars: a, b = 5 urn; c = 1 urn; d, e = 10 urn.

61
 DIATOMS

Figure 2.28. Corethron inerme. a, LMChain of vegetative cells, b. SEM Junction of hooked
spines to sockets of upper valve, c, SEM
2 hooked spines of upper valve of terminal cell. d.
SEM Sockets on valve margin into which spines are inserted, e, SEM Early form of barbed
spines still connecting 2 adjacent cells shortly after cell division, f. SEM Parallel rows of
half girdle bands. Scale bars: a = 10 um; b = 5 urn; c-f =1 urn.

62
 DIATOMS

Figure 2.29. Corethron pennatum. a, LM Vegetative cell, b, SEM Upper valve with ring of
hooked spines; most barbed spines have broken off. c, SEM Junction of hooked spines to
sockets of upper valve, d, SEM Terminal ends of hooked spines, e, SEM Lower valve with
barbed spines and terminal corona of hairs, f, SEM Alternating ligulate girdle bands. Scale
bars: a = 100 urn; b, e, f = 10 jim; c, d =
1 um.

63
 DIATOMS

Figure 2.30. Leptocylindrus mediterraneus a, LM Dividing cell, b, LM Single cell, c,

. SEM
Collapsed cell showing valve face and copulae. d, SEM Detail of valve face, e, SEM
Vegetative cell, view of copulae. Scale bars: a-c = 10 urn; d, e = 2 urn.

64
 DIATOMS

terminal cell bearing 2 types of spines around margin (one long and barbed, the other short
and hooked); hook with a distinct terminal projection at right-angles to the spine and 3
thinner projections below and closely appressed to the spine (Fig. 2.28b). Lower valve with a
marginal ring of barbed spines; spines T-shaped at their base, inserted into sockets on valve
margin. Prior to cell division, spines held within the parent cingulum (girdle) all point in the
same direction; cells free of cingulum have barbed spines orientated downwards and hooked
spines orientated upwards. Cingulum composed of half-bands (Fig. 2.28f)-

distribution: Near Davis Station, East Antarctica; Antarctic inshore waters (Hasle, 1969);
South Georgia and Enderby Land (Hendey, 1937); Antarctic circumpolar distribution
(Thomas & Bonham, 1990).

Corethron pennatum (Grunow) Ostenf. (Ostenfeld, 1903: 6)

BASIONYM: Actiniscus pennatus Grunow, in van Heurck (1883: pi. 82bis, figs 11, 12)

SYNONYMS: Corethron hispidum Castrac. (Castracane, 1886: 86, pi. 20, figs 3, 5)

Corethron criophilum Castrac. (Castracane, 1886: 85, pi. 21, fig. 14)

Corethron murrayanum Castrac. (Castracane, 1886: 86, pi. 21, fig. 4)

Corethron pelagicum Brun (1891: 20, pi. 19, fig. 6)

Corethron valdiviae G.Karst. (Karsten, 1904: 544, tab. 23; 1905: 101, pi. 12, figs 1-10)

ADDITIONAL REFERENCES: Hendey (1937: 325-329, pi. vii, viii), Manguin (1954: 16), Hustedt
(1958: 130), Hasle (1969: 77, 79, figs 23, 42), Fryxell & Hasle (1971: 335, figs 1-6), Priddle &
Fryxell (1985: 54, figs A, C-I), Thomas & Bonham (1990: 105, pi. 12, figs 1, 3, 5), Moisan &
Fryxell (1993: 493), Hasle & Syvertsen (1997: 96, pi. 15), Crawford et al. (1998: 5, figs 1, 6-25)

Fig. 2.29a-f

description: Cells usually solitary, strongly silicified, cylindrical, heterovalvate; pervalvar

axis 20-240 urn. Valves convex, circular; apical axis 5-82 urn. Upper valve with 2 types of
spines around the margin (one long and barbed, the other short and hooked); hook with 2
distinctprojections at right angles to spine (Fig. 2.29e). Lower valve with a marginal ring of
barbed spines (Fig. 2.29c); spines T-shaped at their base, inserted into sockets on valve margin.
Prior to cell division, spines held within the parent cingulum (girdle) all point in the same
direction; cells free of the cingulum have barbed spines orientated downwards and hooked
spines orientated upwards. Spines entirely absent in some specimens. Terminal corona of hairs
sometimes present (Fig. 2.29c). Cingulum composed of alternating open ligulate bands (Fig.
2.29f). Auxospores illustrated by Karsten (1904, tab. 23, as C. valdiviae).

distribution: Antarctic inshore waters (Fryxell& Hasle, 1971); near Davis Station, East
Antarctica (Archer et al, 1996); Heard
(Manguin, 1954); Ross Sea (Hendey, 1937;
I.

Andreoli et al., 1995); KERFIX, 50°40'S 68°25'E, Southern Ocean (Kopczyhska et al,
1998); Southern Ocean, south of the Antarctic Polar Front, cosmopolitan through tropical
waters to the Arctic (Thomas & Bonham, 1990); Weddell-Scotia Sea (Garrison et al., 1987).

Leptocylindrus danicus Cleve (1889: 54)

ADDITIONAL REFERENCES: Cupp (1943: 78, fig. 39), Hasle (1975: 125, fig. 143), Hasle &
Syvertsen (1997: 95, pi. 14, tab. 18)

Fig. 2.18i

DESCRIPTION: Cells solitary or united by valve faces into long chains, cylindrical, weakly
silicified, thin-walled and hyaline; pervalvar axis with numerous rhomboidal girdle bands
(not visible by LM). Valves circular; apical axis 5-16 urn. Of the adjacent cells in a chain,
one valve is slightly concave, the other convex. Small broad pointed spines border the zone
between the valve face and mantle. Labiate processes absent. Chloroplasts numerous, small.

distribution: Cosmopolitan, but absent or scarce in the sub-Antarctic and Antarctic (Hasle
& Syvertsen, 1997); Drake Passage (Rivera, 1983).

65
 DIATOMS

Leptocylindrus mediterraneus (Perag.) Hasle (1975: 124, figs 121-130)

BASIONYM: Lauderia medherranea Perag. (Peragallo, 1888: 89, pi. 6, fig. 45)

SYNONYMS: Dactyliosolen mediterraneus (Perag.) Perag. (Peragallo, 1892: 104, pi. 33, figs 8, 9)

Dactyliosolen mediterraneus var. tenuis Cleve (1897: 300, pi. 1, fig. 14)

Dactyliosolen tenuis (Cleve) Gran (1902: 172)

Dactyliosolen meleagris G.Karst. (Karsten, 1906: 160, pi. 29, fig. 2)

ADDITIONAL REFERENCES: Cupp (1943: 77, fig. 38), Priddle & Fryxell (1985: 62, figs A-D),
Moisan & Fryxell (1993: 493), Hasle & Syvertsen (1997: 95, pi. 15a, b)

Fig. 2.30a-e

description: Cells solitary or united by valve faces into long chains, cylindrical, with
coarsely structured and heavily silicified walls. Valves circular, flat; apical axis 7-35 um,
without spines or processes. Girdle bands numerous, 1-5 in 10 um, punctate, the ends of the
bands in a straight line along the long pervalvar axis (Fig. 2.30c). Cells weakly pigmented;
chloroplasts rarely seen.

distribution: Sea-ice, inshore Davis Station, East Antarctica; Weddell Sea (Moisan &
Fryxell, 1993); Weddell-Scotia Confluence (Garrison et al, 1987); Antarctic to the Arctic
(Hasle & Syvertsen, 1997).

Chains of cells have often been illustrated with the epiphytic flagellate Rhizomonas setigera
(Pavill.) D.J.Patt. et al. attached to the girdle band zone (e.g. Hasle & Syvertsen, 1997,
fig. 15a).Observations suggest that the flagellated (or perhaps even amoeboid) stage emerges
from the central area of the cell and covers frustules in large numbers in the spring. Rings of
four flagellates on separate thecae could represent a later gametic stage. Further study is
required, but it appears that R. setigera is a stage in the life cycle of L. mediterraneus rather
than a distinct epiphyte. If so, the removal of L. mediterraneus from Dactyliosolen by Hasle
(1975) is upheld, especially when one considers the life history of Dactyliosolen blavyanus
(Perag.) Hasle as shown by von Stosch (1985).

Family Melosiraceae

MelosiraadeliaeManguin(1957: 112, pi. 1, fig. 1; 133, pi. 7, fig. 48a, b)

ADDITIONAL REFERENCES: Manguin (1960: 236, pi. 1, figs 11-13), Everitt & Thomas (1986: 5,
fig. 2C)

Fig. 2.31a-f

description: Cells solitary or in distinctive pairs or triplets, or in chains united by mucilage

pads or by surface spines, lenticular to cylindrical; pervalvar axis 9-32 um. Valves circular,
flat or convex; apical axis 14-32 um, covered in small tapered spines or granules, especially
towards the valve margin (Fig. 2.31c). Areolae loculate, opening externally by simple pores.
Rimoportulae in a ring near the mantle edge (Fig. 2.3 le). Girdle bands split, ligulate, with
rows of small pores (Fig. 2.3 le). Chloroplasts small, discoid, numerous.

distribution: Type locality, Cape Margerie, Terre Adelie (Manguin, 1957); coastal waters
off Davis Station, East Antarctica (Everitt & Thomas, 1986).

Specimens collected in the present study were usually smaller (apical axis 15-24 um,
pervalvar axis 9-25 um) than those described by Manguin (14—22 jam and 22-32 um,
respectively).

Melosira sol (Ehrenb.) Kutz. (Kiitzing, 1849: 31)

BASIONYM: Gallionella sol Ehrenb. (Ehrenberg, 1844b: 202, pi. 35, A, 23, fig. 12)

SYNONYMS: Gallionella occulta Ehrenb. (Ehrenberg, 1844b: 202)

Melosira occulus (Ehrenb.) Kutz. (Kiitzing, 1849: 31)
Melosira radiata Grunow (1870: 27)

66
 DIATOMS

Figure 2.31. Melosira adeliae. a, LM Vegetative cell, valve view, b, SEM Valve face
showing surface spines and granules, c, SEM Valve face showing surface spines and
granules, d, SEM Internal surface of valve showing detail of loculate areolae, visible along
fractured edge, e, SEM Girdle view of a solitary cell showing detail of pores in girdle bands
and ring of rimoportulae near valve mantle, f, SEM Cell doublet, recently divided; linking
valve of right hand cell is obscured by partly collapsed girdle bands of the parent cell. Scale
bars: a = 5 urn; b-d = 1 \im; e, f = 5 urn.

67
 DIATOMS

Figure 2.32. Melosira sol. a, LM Chain of 10 cells, with an odd valve at both ends of the
chain, b, LM Surface of linking valve with distinct surface pattern of radial ridges, c, SEM
Chain of 5 cells, oblique view, d, SEM Chain of 3 cells, girdle view, showing small spines
interlinking the 2 cells on the left, and pores and slits on the girdle bands, e, SEM Internal
surface of separation valve, showing radial ridges; note row of external openings of
rimoportulae on mantle, f, SEM Inside of mantle; note the 5 rimoportula apertures near upper
edge. Scale bars: a-d = 10 um; e = 5 urn; f = 1 urn.

68
 DIATOMS

Paraliasol (Ehrenb.) R.M.Crawford (1979a: 209)

ADDITIONAL REFERENCES: Peragallo (1921: 87, pi. 5, figs 6, 7), Manguin (1960: 237, pi. 20, fig.
240), Everitt & Thomas (1986: 9, fig. 3G, as Paralia sol), Thomas & Jiang (1986: 197, as Paralia
sol)

Fig. 2.32a-f

description: Cells in straight chains, cylindrical, linked by marginal spines and interlocking
ridges and grooves on valve surfaces; pervalvar axis 10-38 urn. Valves flat, circular; apical
axis 48-94 um, with Double heterovalvy present; valves of intercalary cells
radial striation.
(i.e. linking or Type 2 valves) with a ring of radial ridges c. midway between centre and
margin (Fig. 2.32e), interlocking with depressions in sibling cells; terminal valves of chain
(i.e. separation or Type 1 valves) with reduced ridges and no marginal spines. Rimoportulae
present on valve mantle (Fig. 2.32e, f). Chloroplasts small, numerous, discoid.

DISTRIBUTION: Plough I., near Davis Station, East Antarctica (Everitt & Thomas, 1986); Cape
Margerie (Manguin, 1960); Petermann I., Booth-Wandel I., Cape Tuxen, King George I.,
Port Lockroy, Argentine Is. (all Antarctic Peninsula) (Peragallo, 1921).

Crawford (1990) remarked that the double heterovalate nature of this genus has lead to
et al.
more species being described in the literature than truly exist. They also suggested that
M. sol, among others, warranted further investigation. Peragallo (1921) observed that
Melosira antarctica van Heurck appears to be a variety of M. sol.

Family Rhizosoleniaceae

Dactyliosolen antarcticus Castrac. (Castracane, 1886: 75, pi. 9, fig. 7)

SYNONYMS: Dactyliosolen antarcticus var. typica Heiden & Kolbe (1928: 509, pi. 8, Fig. 164; fide F.W.Mills,
1934: 574), nom. inval.
Dactyliosolen antarcticus f. typica Heiden & Kolbe (1928: 509, pi. 8, fig. 16), nom. inval.

ADDITIONAL REFERENCES: Hustedt (1927-30: 556, fig. 316), Lebour (1930: 76, fig. 50), Hendey
(1937: 323, pi. 6, fig. 1), Cupp (1943: 76, fig. 37), Manguin (1954: 19), Hustedt (1958: 131),
Manguin (1960: pi. 4, fig. 49; pi. 26, fig. 316), Hendey (1964: 142), Hasle (1975: 118, figs 90-
100, 109-112), Priddle & Fryxell (1985: 58, figs A-E), Moisan & Fryxell (1993: 493), Hasle &
Syvertsen (1997: 166, pi. 32, tab. 37)

Fig. 2.33a-f

DESCRIPTION: Cells solitary or in short chains, cylindrical, straight; pervalvar axis to 140 um.
Valves flat, circular; apical axis 13-90 um, with fine costae radiating from an off-centre (or
even marginal) labiate process (Fig. 2.33b, d). Girdle composed of half-bands, 2-3 in 10 p.m,
coarsely ribbed; ribs 4-15 in 10 |xm, visible by LM (Fig. 2.33a). Each half-band with a
pointed end, the interlocking of the two half-bands forming a distinct oblique line;
interlocking band ends of adjacent band are aligned longitudinally in a straight line along the
pervalvar axis (Fig. 2.33d). Bands with moderately large areolae (Fig. 2.33f); areolae poroid,
elliptical and often flattened at one end, decreasing in size towards end of girdle band,
mostly unoccluded, although fine cribra do occur (Round et al, 1990). Chloroplasts small,
discoid, numerous. Nucleus usually prominent and central in cell.

distribution: Prydz Bay and coastal waters off Davis Station, East Antarctica; an oceanic
species with an almost cosmopolitan distribution (Hendey, 1964); South Atlantic and
Southern Ocean (Hendey, 1937); Liitzow-Holm Bay (Tanimura et al, 1990); possibly
cosmopolitan (Hasle & Syvertsen, 1997).

In D. antarcticus and some specimens of D. tenuijunctus (below), the interlocking girdle

band ends are aligned in a straight line along the pervalvar axis; in other species they are in
an oblique line (Hasle & Syvertsen, 1997).

69
 DIATOMS

Figure 2.33. Dactyliosolen antarcticus. a, LM Vegetative cell, b, SEM Valve and girdle
bands, oblique view; note fine costae radiating from an off-centre labiate process on valve
surface, c, SEM Vegetative cell; SEM Valve and
note off-centre labiate process on valve, d,
girdle bands, oblique view, e, SEM
Coarsely areolated girdle bands of vegetative cell, f,
SEM Girdle bands; note areolae flattened at one end, and pointed ends of each band. Scale
bars: a = 50 um; b, d, f = 5 urn; c, e = 10 um.

70
 DIATOMS

Dactyliosolen tenuijunctus (Manguin) Hasle (1975: 122, figs 1 14-120)

BASIONYM: Rhizosolenia tenuijuncta Manguin (1957: 118, fig. 15; 1960: 270, figs 76, 77)

ADDITIONAL REFERENCES: Priddle & Fryxell (1985: 60, figs A, B), Hasle & Syvertsen (1997: 167,
tab. 37)

Fig. 2.34a-d

description: Cells solitary or in short chains, weakly silicified, cylindrical, straight, slightly
curved or undulating; pervalvar axis 18-55 um. Valves flat, circular to oval; apical axis 6-28
Hin, with fine costae radiating from a central labiate process (Fig. 2.34b). Girdle composed of
half-bands, 5-9 in 10 u.m, finely ribbed, with ribs 28-36 in 10 um; interlocking of the band
ends aligned in a straight or weakly spiral arrangement along pervalvar axis (Fig. 2.34c).
Areolae poroid, rectangular, unoccluded (Fig. 2.34b). Chloroplasts small, discoid, numerous.
Nucleus usually prominent and central in cell.

distribution: Coastal waters near Davis Station, East Antarctica; Drygalski Glacier, Ross
Sea; southern cold-water region (Hasle & Syvertsen, 1997); Terre Adelie (Manguin, 1960);
Weddell-Scotia Confluence (Garrison et al., 1987).

Figure 2.34. Dactyliosolen tenuijunctus. a, SEM Oblique valve view; note off-centre labiate
process, b, SEM Collapsed cell showing ribbed girdle bands, and fine costae on valve, c,
TEM Shadow cast, collapsed cell showing fine costae radiating from labiate process, and
oblique ends to girdle bands, d, TEM Shadow cast, valve internal view with central labiate
process. Scale bars: a, b, d = 1 um; c = 5 um.

71
 DIATOMS

Figure 2.35. Proboscia species, a, P. alata, LM of cell with characteristic slightly curved
proboscis, b-e, P. inermis. b, LM Valve apex in ventral view, c, SEM Proboscis tip of

'winter' form with terminal ring of fine spinulae. d, SEM Valve in lateral view showing
longitudinal slit into which adjacent cell links, e, SEM Shorter proboscis tip of 'spring' form,

with terminal ring of fine spinulae and fine areolae, f, g, P. truncata, SEM Cell apices of
'spring' form in lateral and ventral views. Scale bars: a, b = 10 urn; c-g = 5 urn.

72
 DIATOMS

Guinardia cylindrus (Cleve) Hasle, in Hasle & Syvertsen (1997: 161, pi. 31)

BASIONYM: Rhizosolenia cylindrus Cleve (1897: 24, pi. 2, fig. 12)

SYNONYM: Rhizosolenia antarctica G.Karst. (Karsten, 1905: 95, pi. 11, fig. 1 ; fide Hasle, 1969), Manguin
(1960: 265, pi. 4, figs 56, 57)

ADDITIONAL REFERENCES: Priddle & Fryxell (1985: 84, figs A, B), Priddle et al. (1990: 120,
pi. 15.4, figs 1, 2, as Rhizosolenia cylindrus), Sundstrom (1986: 103, figs 276-278)

Fig. 2.27d, e

description: Cells in long straight chains that break readily, weakly silicified, cylindrical;
pervalvar axis up to 300 um. Valves flat or slightly convex, circular; apical axis 8-50 um;
rows of fine areolae radiating from central external process tube; process hollow, curved to
attach to adjacent cell (Fig. 2.27e); process tube, as well as the impression upon the adjacent
cell into which the tube fits, both visible by LM. Girdle composed of numerous open (split)
bands with ligulae and antiligulae, with fine poroid areolae in a rectangular arrangement.
Chloroplasts numerous, stellate.

distribution: Prydz Bay, coastal waters near Davis Station, East Antarctica; Terre Adelie
(Manguin, 1960, as R. antarctica); Heard I. (Manguin, 1954, as R. antarctica); Weddell-
Scotia Confluence (Garrison et al., 1987).

The closely related Guinardia flaccida (Castrac.) H.Perag. has a distribution that is
"cosmopolitan, but not seen in polar waters" (Hasle & Syvertsen, 1997). Our most southerly
record of this species was at44°09.51'S 147°13.83'E.

Guinardia tubiformis (Hasle) Hasle, in Hasle & Syvertsen (1997: 165, tab. 36)
BASIONYM: Rhizosolenia tubiformis Hasle (1975: 1 15, text fig. 1, fig. 80)

additional reference: Sundstrom (1986: 104, figs 282-285)

description: [after Hasle &

Syvertsen (1997)] Cells in straight close-set chains, fragile and
weakly pervalvar axis 50-100 um. Valves circular, flat, with a single
silicified, cylindrical;
marginal process as a short stout tube; apical axis 3-10 um. Girdle bands distinct, numerous,
split, with ligulae. Chloroplasts small, numerous, peripheral.

distribution: Type locality, 58°00'S 150°05 W, BRATEGG Station 28, South Pacific
,

Ocean (Hasle, 1975); southern cold-water region (Hasle & Syvertsen, 1997).
We have not seen this species.

Proboscia
The most useful feature for distinguishing Proboscia from closely related genera is the
presence of a proboscis: a long, tubular extension of the valve. This is a stouter projection
than the 'process' (rimoportula, labiate process) of Rhizosolenia.

Five species are currently recognised (see discussion in Hasle & Syvertsen, 1997), although
further EM investigations may see taxa of the closely related Rhizosolenia being transferred
to Proboscia. Three species have been reported from Antarctic waters, P. alata, P. inermis
and P. truncata (Jordan et al., 1991). In the same publication, the authors identified 'winter'
and 'spring' forms of these diatoms, noting that winter forms have a longer proboscis.

Proboscia alata (Brightw.) Sundstrom (1986: 99, figs 258-266)

BASIONYM: Rhizosolenia alata Brightw. (Brightwell, 1858: 96, pi. 5, figs 8, 8a)

SYNONYMS: Rhizosolenia alata f. genuina Gran (1908: 56, fig. 68b, c), nom. inval.

Rhizosolenia obtusa auct. non Hensen (1887), sensu Ostenfeld (1908: 56, fig. 69)
See Jordan el al. (1991) for full synonymy.

additional REFERENCES: Hustedt (1927-30: 600, fig. 344), Hendey (1937: 310), Manguin
pi. 26, fig. 318), Hendey (1964: 146, pi.
(1960: 264, 2, fig. 2), Priddle & Fryxell (1985: 76, figs

73
 DIATOMS

A, B), Ferreyra & Ferrario (1983a: 1, pi. 1, 2), Jordan et al. (1991: 65, figs 1-9), Hasle &
Syvertsen(1997: 159, pi. 30)

Fig. 2.35a

description: Cells solitary or in short chains of c. 4 cells, long, cylindrical; apical axis 2.5-
13 pm. Valves subconical, asymmetrical, tapering to an eccentric slightly curved proboscis;
proboscis tip truncate, bearing a short longitudinal slit, a terminal ring of fine spinules and
another small groove; contiguous area developed as a groove into which distal end of sister
cell links; claspers unequal, slightly protruding. Valve areolae fine, poroid, on basal part of
valve. Girdle bands in 2 dorsiventral columns, with pores and loculate areolae. Auxospores
terminal in chain (Hasle & Syvertsen, 1997). Chloroplasts almost certainly discoid (Round et
al, 1990).

distribution: Heard I. and ADBEX; widely distributed in the Antarctic (Jordan et al,
1991); Southern Seas (Hendey, 1937).

Proboscia inermis (Castrac.) R.W.Jord. & Ligowski, in Jordan et al (1991: 66, figs 10-18)

BASIONYM: Rhizosolenia inermis Castrac. (Castracane, 1886: 71, pi. 24, figs 7, 8, [10], 13), De Toni (1894: 830)
SYNONYMS: Rhizosolenia obtusa auct. non Hensen (1887), sensu Hardy & Gunther (1935: 53, fig. 23)
Rhizosolenia inermis f. castracanei Heiden & Kolbe (1928: 521)
Proboscia alata f. inermis (Castrac.) Hust. (Hustedt, 1927-30: 602, fig. 348), sensu Hendey (1937: 311,
"inermis" phase)
Proboscia (ex Rhizosolenia) "alata" sensu Mullins & Priddle (1987: 57, 59)

additional REFERENCES: Cupp (1943: 90, fig. 52E), Ferreyra & Ferrario (1983a, pi. 1, figs 6,
13), Priddle & Fryxell (1985: 76, figs C, D, M, as R. alata), Priddle et al. (1990: 119, pi. 15.3,
figs 6a, b, as R. inermis), Moisan & Fryxell (1993: 293)

Fig. 2.35b-e

DESCRIPTION: Cells solitary or often in chains to long, cylindrical; apical axis 10-19 pm.1 mm
Valves conical, tapering to a stout proboscis; proboscis slightly expanded, with a short
longitudinal slit just below the tip which is surrounded by a ring of fine spinules (Fig. 2.35c);
contiguous area developed as a groove (Fig. 2.35c, e); claspers equal, protruding distally.
Girdle bands in 2 dorsiventral columns.

distribution: Originally described from Antarctic waters (Castracane, 1886); Antarctic

(Priddle et al, 1990).

Proboscia truncata (G.Karst.) Nothig & Ligowski, in Jordan et al (1991: 68, figs 19-29)
BASIONYM: Rhizosolenia truncata G.Karst. (Karsten, 1905: 97, pi. 10, figs 3, 3a)

SYNONYMS: Rhizosolenia inermis f. rostrata Heiden & Kolbe (1928: 522, pi. 8, fig. 162; pi. 9, fig. 167)

Rhizosolenia rostrata (Heiden & Kolbe) Bunt & E.J. F.Wood (1963: 1255)
Rhizosolenia alata f. curvirostris auct. non Gran (1900: 120, pi. 9, fig. 122), sensu Hasle (1969: 142)

Rhizosolenia alata f. truncata (G.Karst.) Hasle (1969: 142)

Rhizosolenia alata auct. non Brightw. (Brightwell, 1858: 56, pi. 5, fig. 8), sensu Fenner et al. (1976: pi. 13,
fig. 1)

Rhizosolenia alata f. indica Nothig, in Nothig & Nothig (1988: 74, fig. 42a, c)

ADDITIONAL REFERENCE: Priddle et al. (1990: 119, pi. 15.4, fig. 4)

Fig. 2.35f, g
DESCRIPTION: Cells solitary or often cells; individual cells 140-235 pm
in long chains of up to 8
long, cylindrical; apical axis 16-30 pm
Valves dissimilar; one with a short
diameter.
proboscis, the other with a long one. Short proboscis truncate, angled at c. 45° to pervalvar
axis (lateral view), with spinules and a longitudinal slit at tip; claspers equal, protruding.
Long proboscis tapered, straight or slightly angled (lateral view), slightly wider, with
spinules and a short longitudinal slit just below tip; claspers, when present, equal,
protruding. Girdle bands not distinguishable.

74
 DIATOMS

—a -b

'-h

Figure 2.36. Rhizosolenia species, a, R. polydactyla f. polydactyla, LM of cell apex, with

small rounded otaria. b-f, R. antennata f. semispina. b, LM Apex of cell with small pointed
otaria. c, SEM Solitary cell, d, SEM Apices on interconnected cells, e, SEM Cell valve in
ventral view; otaria and claspers visible, f. SEM Cell valve in dorsal view; otaria visible, g,
h, R. antennata f. antennata. g, LM Apex of cell, h, LM Apex of cell. Scale bars: a, b, d, e,

g, b = 10 urn; c = 100 urn; f =1 urn.

75
 DIATOMS

Figure 2.37. Rhizosolenia species, a, R. crassa, LM Valve apex (from Armand & Zielinski.
2001). b, R. curvata, LM Valve apex, c, R. hebetata f. semispina, LM Valve apex. d. R.
polydactyla f. polydactyla, LM Valve apex, e, R. sima f. sima, LM Valve apex; note broad
otaria (from Armand & Zielinski, 2001). f. R. styliformis, LM Valve apex, (from Armand &
Zielinski, 2001). Scale bars: a-f = 10 urn.

76
 DIATOMS

distribution: Prydz Bay, near Davis Station, East Antactica; Drygalski Glacier, Ross Sea;
recorded only from Antarctic waters (Jordan et al., 1991); Antarctic Peninsula (Ferreyra &
Ferrario, 1983a).

Ferreyra & Ferrario (1983a) suggested that the long proboscis type is a winter form, and this
was confirmed by Jordan et al. (1991).

Rhizosolenia

Species of Rhizosolenia have long cylindrical cells with numerous girdle bands. They are
readily distinguished by having a single spine or process (also called a rimoportula or labiate
process) arising from each valve with a groove on each to accommodate the process of the
adjacent valve (Round et al, 1990). Eleven species or forms of Rhizosolenia representing
two sections are generally recognised in the Antarctic (Priddle et al., 1990; Jordan et al.,

1991). In their study of rhizosolenioid remains in Southern Ocean sediments, Armand &
Zielinski (2001) recognised of the species groupings defined by Priddle et al. (1990) and
all

an additional form with affinities to R. hebetata f. semispina. Rhizosolenia chunii is

referrable to section Imbricatae; all other taxa belong to section Rhizosolenia.

Rhizosolenia acuminata (H.Perag.) H.Perag., reported rarely from Antarctic waters, is

insufficiently known to warrant a full description here.

Rhizosolenia antennata (Ehrenb.) N.E.Br, f. semispina Sundstrom (1986: 44, figs 20,
114, 116-118)

ADDITIONAL REFERENCES: Priddle et al. (1990: 1 17, pi. 15.1, fig. 6), Kang & Lee (1995: 260, fig.

6D), Hasle & Syvertsen (1997: 149, tab. 33), Armand & Zielinski (2001, pi. 3a, figs 4, 5)

Fig. 2.36b-f

description: Cells solitary, in pairs or in long chains, long, cylindrical or slightly swollen.
Valves acutely conical, bilaterally symmetrical; apical axis 6.5-42 um. Process long (often
broken during the preparation of material for microscopy), tapering. Otaria visible by LM,
pointed, extending for several microns along the basal part of the process. Claspers present,
visible by LM.

distribution: Heard I.; inshore waters of Davis Station, East Antarctica; Southern Ocean,
offshore Prydz Bay, 67°01'S 77°43'E; Bransfield Strait (Kang & Lee, 1995); Weddell and
Bellingshausen Seas (Hendey, 1937); surface sediments from Weddell Sea and Scotia Sea
(Armand & Zielinski, 2001).

Rhizosolenia antennata f. semispina possesses otaria. There has been considerable confusion
between this taxon and R. hebetata f. semispina, and both Sundstrom (1986) and Priddle et
al. (1990) suggested that many early records of the latter actually represent R. antennata

f. semispina. Armand & Zielinski (2001) further suggested that if these are conspecific, then

R. antennata f. semispina is one of the most common Rhizosoleniae in the Southern Ocean.

Rhizosolenia antennataf. antennata (Fig. 2.36g, h) is thought to represent the resting spore of

R. antennata semispina (Sundstrom, 1986; Priddle et al, 1990; Hasle & Syvertsen, 1997).
f.

The former has two (paired) processes arising from each valve, the otaria are contiguous and
claspers are lacking. Its distribution is generally assumed to include Antarctic coasts,
northwards to the edge of the sea-ice zone. Reports from Antarctic waters have appeared under
the names R. bidens (Kopczyhska et al., 1986) and R. hebetata f. bidens (Priddle et al, 1990).

Rhizosolenia chunii G.Karst. (Karsten, 1905: 99, pi. 11, fig. 5)

SYNONYM: 1 Rhizosolenia murrayana Castrac. (Castracane, 1886: 72, pi. 24, fig. 12)

ADDITIONAL REFERENCES: Hendey (1937: 313), Hustedt (1958: 133), Manguin (1960: 266, pi. 4,

fig. 58a, b, 59, 60; pi. 26, figs 319-321; pi. 27, fig. 322a, b), Sournia et al. (1979: 191, figs 17,
29), Priddle & Fryxell (1985: 78, figs A-E), Priddle et al. (1990: 118, pi. 15.3, fig. 2), Moisan &
Fryxell (1993: 293). See Armand & Zielinksi (2001) for additional references.

77
 DIATOMS

description: Cells in short chains, weakly silicified, cylindrical, almost rectangular in girdle
view; pervalvar axis 47-1 12 urn (fide Manguin, 1960). Valves flat or slightly convex; apical
axis 20-28 urn (26-32 um fide Hendey, 1937; 12-19 [im fide Manguin, 1960). Process
arising from the valve margin, short, stout, hollow at the base. Otaria lacking. Intercalary
girdle bands 2-3 in 10 um, arranged in 2 lateral (cf. dorsiventral) columns forming zig-zag
rows; bands finely striate; striae 18-23 in 10 um. Chloroplasts small, numerous.

distribution: Weddell-Scotia Confluence (Garrison et al., 1987); Bellingshausen Sea

(Hendey, 1937; Ligowski, 1993); Terre Adelie (Manguin, 1960); sub-Antarctic (Hasle, 1969);
Antarctic (Sundstrom, 1986).

Rhizosolenia chunii poorly known from modern Antarctic studies. Sundstrom (1986) noted
is

that this species stands out because of its "short" pervalvar axis. He also indicated that re-
circumscription was warranted due to the conspecificity with Castracane's R. murrayana and
the incomplete observations he (Castracane) had made. Sediment records are limited, and the
species requires further study (Armand & Zielinski, 2001).

Rhizosolenia crassa A.F.W.Schimp., in Karsten (1905: 99, pi. 11, fig. 6)

ADDITIONAL REFERENCES: Hendey (1937: 314), Priddle & Fryxell (1985: 80, figs A, B), Sundstrom
(1986:59, figs 26-27, 139-143), Priddle et al. (1990: 118, pi. 15.3, fig. 1), Hasle & Syvertsen (1997:
150, pi. 28, tab. 34), Armand & Zielinski (2001: pi. 3 A, figs 10, 1
1)

Fig. 2.37a

description: Cells solitary or in pairs or short chains, large, cylindrical. Valves short-conical,
bilaterally symmetrical, finely areolate; areolae 27 in 10 um; apical axis 100-165 um (160—
220 um, fide Hendey, 1937). Process c. 50 um long, robust basally, tapering distally (often
broken during the preparation of material for microscopy). Otaria present, ending at the
junction of the valve apex and the basal part of the process; distal margin of otaria concave.
Claspers prominent. Intercalary girdle bands in 2 and 4 (6, fide Karsten, 1905: pi. 1 1, fig. 6a)
dorsiventral columns, finely areolate; areolae 22-24 in 10 um. Chloroplasts small, numerous.

distribution: Southern cold-water regions (Hasle & Syvertsen, 1997); Weddell Sea, South
Sandwich Is. and South Georgia (Hendey, 1937); Wilkes Coast, 62°S 135°E and 64°S 115°E
(Wood, 1960); Antarctic Permanent Open-Ocean Zone to Sea-Ice Zone (Armand & Zielinski,
2001).

Rhizosolenia curvata O.Zacharias (1905: 120, text fig.)

SYNONYM: Rhizosolenia curva G.Karst. (Karsten, 1905: 97, pi. 11, fig. 2)

ADDITIONAL REFERENCES: Hendey (1937: 314), Hustedt (1958: 133), Manguin (1960: 268, pi. 5,

figs 64-66), Priddle & Fryxell (1985: 82, figs A-F), Priddle et al. (1990: 118, pi. 15.2, fig. 3),
Hasle & Syvertsen (1997: 147, pi. 26, tab. 32), Armand & Zielinski (2001: pi. 3A, figs 1-3)

Fig. 2.37b

DESCRIPTION: Cells solitary or paired, long, cylindrical, distinctly curved; pervalvar axis 10-
95 um long (or longer fide Manguin, 1960). Valves conical, bilaterally symmetrical; apical
axis 20-135 um. Process long, uniformly tapering (often broken during the preparation of
material for microscopy). Otaria small, rounded, arising on the distal part of the valve (rather
than the process). Contiguous area with distinct marginal ridges. Claspers small. Intercalary
girdle bands arranged in 2 dorsiventral columns; bands 20 in 10 um, finely areolate.

distribution: Terre Adelie (Manguin, 1960); South Georgia and Drake Passage (Hendey,
1937); sub- Antarctic/Polar Front (Hasle & Syvertsen, 1997); Southern Ocean (Hart, 1937).

Armand & Zielinski (2001) summarised the distribution of/?, curvata as sub-Antarctic,
limited southward by the position of the Polar Front.

78
 DIATOMS

Figure 2.38. Rhizosolenia imbricata. a, b, LM Single cell, c, SEM Single cell, d, SEM Cell
apex showing flattened valve, e, LM Cell apex, ventral view with swollen base of process.
Scale bars: a-c = 10 um; d, e = 1 um.

79
 DIATOMS

Figure 2.39. Rhizosolenia simplex, a, LM Valve with spine, b, LM Valve with spine, c, SEM
Valves from ajoining cells, still linked together, d. SEM Valve with spine. Scale bars: a-d =
10 ^irn.

80
 DIATOMS

Rhizosolenia hebetata (Bailey) Gran f. semispina (Hensen) Gran (1904: 527, pi. 27,
figs 11, 12)

BASIONYM: Rhizosolenia semispina Hensen (1887: 84, pi. 5, fig. 39)

SYNONYM: Rhizosolenia styliformis var. semispina (Hensen) G.Karst. (Karsten, 1905: 96, pi. 10, figs 4, 4a,
4b), sensu Wimpenny (1946: 277, text fig. Hi, i)

ADDITIONAL REFERENCES: Gran (1905: fig. 67b; 1908: 55, fig. 67b), Cupp (1943, fig. 50B),
Manguin (1960: 268, pi. 5, figs 67, 68; pi. 27, figs 323, 324), Fenner et al. (1976, fig. 6), Priddle
& Fryxell (1985: 86, figs A-H), Sundstrom (1986: 48, figs 19, 20, 114, 116-118), Priddle et al.
(1990: 117, pi. 15.1, fig. 4), Tanimura (1992: fig. 3.15), Moisan & Fryxell (1993: 293), Hasle &
Syvertsen (1997: 149, pi. 21 pro parte)

Fig. 2.37c

description: Cells solitary or in pairs or chains, long, cylindrical; pervalvar axis 350-775
Urn. Valves acutely conical, bilaterally symmetrical, finely areolate; areolae 22-24 in 10 urn.
Apical axis 4.2-25 (im (5-12 um fide Cupp). Process long (often broken during the
preparation of material for microscopy), tapering. Otaria and claspers lacking. Intercalary
girdle bands finely areolate; areolae 20-23 in 10 urn. Chloroplasts small, numerous.

distribution: Weddell Sea (Moisan & Fryxell, 1993); Weddell-Scotia Confluence (Garrison
etal, 1987).

As a result of the confusion between R. hebetata f. semispina, R. antennata f. semispina and

R. styliformis, the distribution pattern of R. hebetata f. semispina in Antarctic surface waters

is unclear and must be treated with extreme caution (Armand & Zielinski, 2001).

Sundstrom (1986) emended the description of R. hebetata f. semispina (otaria-bearing) and

R. hebetata f. hebetata (otaria-lacking), and limited their distribution to the Northern
Hemisphere. Armand & Zielinski (2001) discussed in detail the confusion over the
identification of Southern Ocean Rhizosolenia species in the light of the various erroneous
early descriptions.

Some records of/?, hebetata f. semispina from the Antarctic (e.g. Hendey, 1937; Hustedt,
1958) possibly refer to R. styliformis (see Priddle et al., 1990). Armand & Zielinski (2001)
also noted that "many workers found it difficult to separate Rh. styliformis from Rh. hebetata
f. semispina (e.g. Hart 1934, 1942, Hargraves 1968, Hasle 1969, Simonsen 1974, Fenner et

al. 1976)".

Rhizosolenia hebetata f. hebetata is now considered to be the probable resting spore of R.

hebetata f semispina (Priddle et al., 1990). It is unlikely that either form exists in the
Southern Ocean.

Rhizosolenia imbricata Brightw. (Brightwell, 1858: 95, pi. 5, fig. 6)

SYNONYMS: Rhizosolenia striata Grev. (Greville, 1865: 234, pi. 3, fig. 4)

Rhizosolenia shrubsolei Cleve (1881: 26)

Rhizosolenia imbricata var. striata (Grev.) Grunow, in van Heurck (1882: pi. 79, figs 3, 7)

Rhizosolenia imbricata var. shrubsolei (Cleve) Schroder (1906b: 346)

ADDITIONAL REFERENCES: Hustedt (1927-30: 580, fig. 331), Hendey (1937: 316; 1964: 149, pi. 3,
fig. 1), Priddle & Fryxell (1985: 90, figs A-E, as R. shrubsolei), Sundstrom (1986: 80, figs 200-

208), Hasle & Syvertsen (1997: 155, pi. 29, tab. 35)

Fig. 2.38a-e

description: Cells solitary or in pairs or chains, long, cylindrical; pervalvar axis to 400 um.
Valves obliquely conical to flattened, finely areolate; apical axis 2.5-57 um. Process short,
8-18 um, swollen at the base and narrowing abruptly to a distal tube (Fig. 2.38d). Otaria
small, extending along the swollen part of the process. Claspers conspicuous. Girdle bands
forming 2 dorsiventral columns of segments, finely areolate; areolae 10-18 in 10 um,
arranged in oblique striae.

81
 DIATOMS

distribution: Prydz Bay, ADBEX; Southern Ocean, south of Australia; Weddell-Scotia

Confluence (Garrison et al., 1987, as R. schruhsolei); Southern Ocean (Priddle & Fryxell,
1985).

Although agreeing with other morphological features described for R. imbricata, cells
Ocean (Thomas, ADBEX) have a shorter valve process (2.8-3.2
collected from the Southern
urn long).

Rhizosolenia polydactyla Castrac. (Castracane, 1886: 71, pi. 74, fig. 2) f. polydactyla
SYNONYMS: Rhizosolenia styliformis var. oceanica Wimpenny (1946: 279, text fig. Id, e)

Rhizosolenia styliformis auct. non Brightwell (1858), sensu Hendey (1937: pi. 11, figs 15-17)
Rhizosolenia styliformis var. longispina auct. non Hustedt (1934), sensu Okuno (1952, pi. 2, fig. 1)

ADDITIONAL REFERENCES: Sundstrom (1986: 24, figs 7, 70-72, 74, 76, 77, 79), Priddle et al.

(1990: 118, pi. 15.2, figs 4, 5), Hasle & Syvertsen (1997: 150, pi. 27, tab. 33), Armand &
Zielinski (2001: pi. 3A, figs 6-9, as f. polydactyla; pi. 3B, figs 2, 3, as f. squamosa)

Figs 2.36a, 2.37d

description: Cells usually solitary, long, cylindrical. Valves conical, bilaterally

symmetrical; apical axis 15-105 um (56-105 um in f. squamosa); areolae 26-28 in 10 urn.
Process 18-34 um long, tapering distally from c. midway along its length. Otaria large,
rounded, ending clearly above the base of the process and at right-angles. Contiguous area
and claspers conspicuous. Girdle bands forming 2 dorsiventral columns of segments,
areolate; areolae 20-23 in 10 um.

distribution: Southern cold-water regions (Hasle & Syvertsen, 1997); Marguerite Bay,
Bellingshausen Sea and Amundsen Sea (Mangin, 1915); Bellingshausen Sea, South Georgia,
Drake Passage (Hendey, 1937).

Based on their sediment distribution observations, Armand & Zielinski (2001) suggested this
is the most common open-ocean species of Rhizosolenia.

Rhizosolenia polydactyla f. squamosa Sundstrom is generally regarded as the otaria-lacking

resting spore of R. polydactyla f. polydactyla. However, the girdle bands off. squamosa form
several pairs of segment columns, and the valves are of a coarser structure than in
f. polydactyla. While the former lacks otaria, the rimportula is heavily silicified. An
intermediate form was illustrated by Priddle et al. (1990: pi. 15.2, fig. 6). Sundstrom (1986)
listed R. styliformis var. latissima Brightw. ex Freng. & Orlando (Frenguelli & Orlando,
1958: 17, fig. 6) as a synonym of R. polydactyla f. squamosa.

Rhizosolenia rhombus G.Karst. (Karsten, 1905: 97, pi. 10, fig. 6a-c)

ADDITIONAL REFERENCES: Hendey (1937: 317), Hustedt (1958: 134), Priddle & Fryxell (1985: 86,
figs A-D), Priddle et al. 990: 20, pi. 5.2, fig. 7)
( 1 1 1

description: Cells solitary, large, rhombic in girdle view. Valves moderately convex to flat,
bearing a stout process displaced to one side; apical axis 10-280 um. Otaria large, rounded, 1

extending for most of the length of the process (Karsten, 1905). Contiguous area
conspicuous. Girdle bands forming 2 dorsiventral columns of segments, with areolae
arranged in quincunx (4 at the corners of a square and one at the centre; Hendey, 1937).

DISTRIBUTION: Antarctic (Priddle et al., 1990); South Georgia (Hendey, 1937).

Priddle et al. (1990: 1 15) indicated that this species should be transferred to a genus yet to be
described.

Rhizosolenia sima Castrac. (Castracane, 1886: 71, pi. 24, fig. 20) f. sima
ADDITIONAL REFERENCES: Sundstrom (1986: 62, figs 28, 144-149), Priddle et al. (1990: 118, pi.

15.2, fig. 1, as f. sima; fig. 2, as f. silicea); Hasle & Syvertsen (1997: 150, pi. 27, tab. 33),
Armand & Zielinski (2001 : fig. 3L, as R. sima f. sima; fig. 13a, b, as R. sima f. silicea)

82
 DIATOMS

Fig. 2.37e

description: Cells usually solitary or in pairs, long, cylindrical, straight or slightly curved.
Valves conical, bilaterally symmetrical; apical axis 12-39 urn (18-40 pm in f. silicea).
Process bulbous, tapering abruptly to a thin-walled distal tube. Otaria large, rounded, arising
on the valve apex and extending along the basal part of the process, also extending laterally.
Contiguous area only distinct near the sister-cell process. Claspers conspicuous. Girdle bands
forming 2 dorsiventral columns of segments, areolate; areolae 31-36 in 10 urn.

distribution: Southern Ocean, south of Australia; Admiralty Bay, Bellingshausen and

Weddell Seas (Ligowski, 1993).

Rhizosolenia sima f. silicea Sundstrom (1986: 63, figs 29, 150-154), also recorded from
southern cold-water regions, is generally regarded as the heavily silicified resting spore off.
sima. Forma silicea lacks otaria, claspers and a contiguous area, and has a robust process
arising from a thickened valve apex (Hasle & Syvertsen, 1997).

Rhizosolenia simplex G.Karst. (Karsten, 1905: 95, pi. 10, fig. 1)

SYNONYMY: probably conspecific with R. torpedo G.Karst. (Karsten, 1905: 95, fig. 2a)

ADDITIONAL REFERENCES: Manguin (1954: 21), Hustedt (1958: 134), Priddle & Fryxell (1985: 92,
figs A-D), Sundstrom (1986), Priddle et al. (1990: 118, pi. 15.3, fig. 3), Armand & Zielinski
(2001: 283, fig. 4K, L)

Fig. 2.39a-d

DESCRIPTION: Cells long, cylindrical, comparatively weakly silicified. Valves conical; apical
axis 5-48 pm. Process uniformly tapering, often conical, 15-20 urn long. Otaria, contiguous
area and claspers absent. Intercalary girdle bands in 2-4 dorsiventral columns, finely
areolate.

distribution: Oceanic sub-Antarctic regions (Hasle, 1969; Semina, 1979); Heard I.

(Manguin, 1954); Bellingshausen and Weddell Seas (Ligowski, 1993).

Rhizosolenia styliformis Brightw. (Brightwell, 1858: 95, fig. 5a, b, d, non 5c)
SYNONYMS: Rhizosolenia styliformis var. polydactyla van Heurck (1909: 28, figs 66, 67, 70, 71 , 74, 75)

Rhizosolenia styliformis var. longispina Hust., in Schmidt (1914: 316, figs 5-8, 12); Sundstrom (1986: 15,
figs 5, 47-56); Everitt & Thomas (1986: 5, fig. 2D)
Rhizosolenia styliformis f. latissima Brightw., in Peragallo (1892), Manguin (1960: 270, pi. 5, fig. 71; pi. 6,

figs 72-75)

ADDITIONAL REFERENCES: Hendey (1937: 320, pi. 11, figs 15-17), Priddle & Fryxell (1985: 94,
figs A-G), Priddle et al. (1990: 117, pi. 15.3, fig. 4), Armand & Zielinski (2001; 261, figs 4a, 5A-
L, from the holotype slide)

Fig. 2.37f

DESCRIPTION: Cells solitary or paired, long, cylindrical. Valves 23-90 um diameter, markedly
asymmetrical, with ventral margin (excluding process) extending for c. half of cell length.
Process 30-50 um long, uniformly tapering; claspers visible. Otaria small, rounded,
terminating below process base or extending 1-2 pm along it. Girdle bands in 2 dorsiventral
rows, the first 2 bands differing from others due to asymmetry of valves.

distribution: Arctic (Priddle et al, 1990); Kerguelen (Hustedt, 1958, as R. styliformis

Bellingshausen Sea and Weddell Sea (Ligowski, 1993).
var. longispina);

The circumscription of R. styliformis has been the subject of much discussion and revision
(e.g. Sundstrom, 1986; Armand & Zielinski, 2001). Confusion regarding its identification has
seen most records of R. styliformis in the pre-2000 literature re-named R. antennata
f. semispina, R. polydactyla or R. curvata. However, R. styliformis is distinguishable from

these taxa by its otaria ending at or below the process base. Researchers are urged to

83
 DIATOMS

compare their specimens with recent photographic images of the holotype slide provided by
Hasle (1975) and Armand & Zielinski (2001) to ensure correct identification.

Family Thalassiosiraceae

Lauderia annulata Cleve (1873: 8, pi. 1, fig. 7)

SYNONYM: Lauderia borealis Gran (1900: 109, pi. 9, figs 1-8)

ADDITIONAL REFERENCES: Hasle & Syvertsen (1996: 36, pi. 1), Round et al. (1990: 150)

Fig. 2.40a-c

description: Cells in chains attached by threads from strutted processes on valve faces,
discoid. Valves circular, 24-75 urn diameter. Areolation appearing as barely discernible ribs
radiating from a central annulus; ribs more than 30 in 10 urn, anastomosing near the margin.
Strutted processes scattered over valve surface and in a conspicuous marginal ring; processes
10 in 10 um, with well-developed external tubes. Labiate process solitary, just inside
marginal ring of strutted processes. Plastids small, discoid.

distribution: Coastal waters off Davis Station, East Antarctica.

Superficially this species resembles members of the genus Porosira, but it has distinctive
external extensions to the strutted processes.

Planktoniella sol (G.C.Wall.) Schutt (1892: 258, fig. 64)

basionym: Coscinodiscus sol G.C.Wall. (Wallich, 1860: 38, pi. 2, fig. 1)

ADDITIONAL REFERENCES: Ricard (1987: figs 124-128), Round et al. (1990: 134), Hasle &
Syvertsen (1997: 40, pi. 2, tab. 4)

Fig. 2.40d, e

DESCRIPTION: Cells solitary, discoid, with a wing of organic extrusions (stiffened mucilage?)
lying around the periphery of the cell. Valves circular, 10-60 um diameter, with a shallow
mantle. Areolation radial to tangential; areolae 5-9 in 10 um, opening internally via fine
cribra and externally by larger foramina. Strutted processes in a single marginal ring;
processes 4-5 in 10 um, as well as single process near valve centre. Two labiate processes
present just inside marginal ring of strutted processes. Plastids small, discoid.

distribution: Southern Ocean, south of Australia.

Porosira glacialis (Grunow) Jorg. (Jorgensen, 1905: 97, pi. 6, fig. 7)

BASIONYM: Podosira hormoides var. glacialis Grunow (1884: 108, pi. E, fig. 32)
SYNONYMS: Podosira glacialis (Grunow) Cleve (1896: 12, pi. 2, figs 17-20)
Lauderia glacialis (Grunow) Gran (1900: 1 1
1, pi. 9, figs 10-14)
Porosira antarctica O.G.Kozlova (1962: 8, pi. 2, fig. 1; pi. 3, fig. 1; 1967: 54, fig. 1)

ADDITIONAL REFERENCES: Manguin (1954: 20), Jouse et al. (1962: 66, pi. 4, fig. 16, as
P. antarctica), Priddle & Fryxell (1985: 142, figs A-C), Hasle & Syvertsen (1997: 43, pi. 3, tab.

5)

Fig. 2.41a-f

DESCRIPTION: Cells solitary or attached by mucilage threads from valve surfaces to form
loose chains (Fig. 2.4 If), discoid to cylindrical. Valves circular, 30-40 um diameter;
pervalvar axis 12-64 um. Areolation radial to irregular, sometimes indistinct; areolae 25-26
in 10 um, loculate, with external foramina and internal cribra. Strutted processes scattered
over valve face (Fig. 2.41c, d), 4-6 in 10 um, simple or with short tubes from which
mucilage threads emerge. Labiate process solitary, inside margin (Fig. 2.41d). Girdle bands
with areolae similar to valves (Fig. 2.41e). Chloroplasts small, discoid, numerous.

84
 DIATOMS

Figure 2.40. Lauderia annulata and Planktoniella sol. a-c, Lauderia annulata. a, SEM
Single cell, valve view of partly collapsed cell, b, SEM Detail of central annulus, and fine
threads from strutted processes, c, SEM Fractured valve margin showing detail of tubular
strutted processes emergent on internal and external valve surfaces, d, e, Planktoniella sol. d,
LM Solitary cell with wing-like extrusion lying around the cell periphery, e, SEM Solitary
cell with wing-like extrusion lying around the cell periphery. Scale bars: a-e = 10 pm.

S5
 DIATOMS

Figure 2.41. Porosira glacialis. a, LM Vegetative cell, b, SEM Valve, internal view, detail
of strutted process and single labiate process, c, SEM Valve, internal view, d, SEM
Vegetative cell, oblique view, e, SEM Valve, internal view; note strutted processes and
single marginal labiate process, f, SEM Vegetative cells joined by mucilage threads to form a
chain. Scale bars: a, f = 10 urn; b, c, e = 5 urn; d =
1 urn.

86
 DIATOMS

Figure 2.42. Porosira pseudodenticulata a, . LM

Vegetative cell, b, SEM Valve, external
view, c, SEM Vegetativeoblique view, d, SEM Vegetative cell with mucilage threads
cell,
emerging from strutted processes on valve face, e, Valve, internal view; note scattered
strutted processes, f, SEM Valve, internal view of strutted processes and single labiate
process, g, SEM External view of labiate process. Scale bars: a-e = 10 urn; f g =
,
1 urn.

87
 DIATOMS

Figure 2.43. Thalassiosira ambigua. a, LM Single cell with distinct chloroplasts. b, SEM
External valve view of 2 cells, c, SEM Girdle view of a collapsed cell, d, SEM Valve view of
cell with strands emanating from marginal processes, e. SEM Valve view of cell. Scale bars:
a. b, d, e = 5 urn; c = 2 urn.
 DIATOMS

distribution: Southern Ocean, south of Australia and near Davis Station, East Antarctica;
type locality, Zemlya Frantsa-Iosifa (Franz Josef Land), Arctic Ocean (Grunow, 1884);
Heard I. (Manguin, 1954); bipolar distribution (Hasle & Syvertsen, 1997).

Cells collected near Davis Station were discoid in shape with a pervalvar axis of 12-22 urn.

Porosira pseudodenticulata (Hust.) Jouse, in Kozlova (1962: 10, fig. 3, no. 2)

BASIONYM: Coscinodiscus pseudodenticulata Hust. (Hustedt, 1958: 1 17, figs 20, 21)

SYNONYM: IPodosira adeliae Manguin (1960: 240, pi. 1, figs 10-12; pi. 21, figs 250, 251)

ADDITIONAL REFERENCES: Jouse (1962: 66, pi.

et al. 4, figs 15, 16), Priddle & Fryxell (1985: 145,
figs A, B), Andreoli et al. (1995: 469, fig. 15), Hasle & Syvertsen (1997: 43, pi. 3, tab. 5)

Fig. 2.42a-g

description: Cells solitary or attached by mucilage threads from the valve surfaces to form
loose chains (Fig. 2.42d), discoid to cylindrical. Valves circular, 36-80 pm diameter;
pervalvar axis 10-30 pm. Areolation radial; striae spiralling; areolae 10-16 in 10 pm,
loculate, with external foramina and internal cribra. Strutted processes scattered over valve
face, although sometimes sparse in the centre (Fig. 2 Aid, e), 3-4 in 10 pm, simple or with
short tubes from which mucilage threads emerge (Fig. 2.42d). Labiate process solitary, inside
margin (Fig. 2.42f). Girdle bands with areolae similar to valves (Fig. 2.42d). Chloroplasts
small, discoid, numerous.

distribution: Sea-ice near Davis Station, East Antarctica, and Southern Ocean, south of
Australia; type locality,South Atlantic Ocean and Antarctic, c. 44-69°S and 0-6°30'E
(Hustedt, 1958); Weddell-Scotia Confluence (Garrison et al., 1987); Antarctic, circumpolar
(Hasle, 1973); Terra Nova Bay, Ross Sea (Andreoli et al., 1995); Liitzow-Holm Bay
(Tanimura et al., 1990).

Specimens from Prydz Bay, photographed here, fit the descriptions of P. pseudodenticulata,
but they are somewhat smaller in size and can have a higher density of areolae (10-16 in
10 pm) than was reported by other authors (10-12 in 10 pm).

Thalassiosira ambigua O.G. Kozlova (1962: 8, pi. 2, fig. 7)

ADDITIONAL REFERENCES: Kozlova (1967: 57, figs 4, 14), Fryxell & Hasle (1983: 57, figs 15-28),
Johansen & Fryxell (1985: 158, figs 24, 36), Fryxell & Johansen (1990: 99, pi. 1 1.9, fig. 10)

Fig. 2.43a-e

DESCRIPTION: Cells solitary or chain-forming by a chitinous strand from central or marginal

strutted process. Valves circular, 6.5-14 pm diameter; pervalvar axis 2-4.6 pm. Areolae
radial, 24-32 in 10 pm in the centre; tangential areolae absent. Strutted processes 1 or 2 in
valve centre, with a single ring of processes at the margin, 6-10 in 10 pm; processes with
external projection. Labiate process solitary, located within the marginal ring.

distribution: Near Davis Station, East Antarctica.

Thalassiosira antarctica Comber (1896: 491, pi. 11)

SYNONYMS: Thalassiosira antarctica G.Karst. (Karsten, 1905: 73, pi. 2, figs 2, 3), nom. inval.
Thalassiosira fallax Meunier (1910: 268, pi. 30, figs 1-4) (see Syvertsen 1979: 59)

Thalassiosira antarctica var. borealis G. A. Fryxell, Doucette & G.F.Hubb. (Fryxell et al., 1981: 323, figs 21-
24, 28-33)

See VanLandingham (1978: 3992) for full synonymy.

ADDITIONAL REFERENCES: Hendey (1937: 237), Manguin (1954: 22), Hasle & Heimdal (1968:
357, figs 1-23), Fryxell et al. (1981: 322-332, figs 1-5, 11-13, 17-20, 26,27, 35A, B), Semina &
Makarova (1983: 613, pi. 2, figs 5, 6; pi. 4, figs 1-6; pi. 5, figs Johansen & Fryxell (1985:
1-6),
158, figs 15-17, 37-39), Fryxell & Johansen (1990: 99, pi. 11.9, figs 1-3), Hasle & Syvertsen

89
 DIATOMS

(1990d: 289, figs 25-27), Moisan & Fryxell (1993: 293), Hasle & Syvertsen (1997: 66, pi. 8, tab.

10)

Fig. 2.44a-d

description: Cells chain-forming, held together by chitinous strands emerging from central
cluster of strutted processes (Fig. 2.44a, d); distancebetween cells usually 2-3 times as great
as pervalvar dimensions of cells. Larger cells less strongly silicified. Valves circular, 14-56
Urn diameter, becoming markedly convex with reduction in size. Areolation radial, bifurcate
to fasciculate; areolae 17-20 in 10 um, loculate, opening externally via a simple foramen,
internally via an evenly pored velum (Fig. 2.44b-d). Strutted processes 2-several, 4- or 5-
sided, operculate (Fig. 2.44c), with small external extensions not visible by LM, 8-16 in 10
um on mantle, 4-14 in 10 um in the centre. Labiate process large, solitary, sessile, located in
outer ring in valves with 1 marginal ring (Fig. 2.44b, d), but located between rings in those
valves with 2 or 3 marginal rings of strutted processes. Internal opening of labiate process
aligned with radius of valve (Fig. 2.44b, d); external opening elongated and similarly
orientated. Cingulum with an areolate valvocopula; copula with a line of pores proximal to
valvocopula and some additional pores on the ligula. Pleurae apparently unstructured, 3 or 4
and spiralling to the right. For a complete description, see Fryxell et al. (1981) and Johansen
& Fryxell (1985).

DISTRIBUTION: Type locality, South Atlantic, in the area of the South Shetland Is.; Southern
Ocean and sub-Antarctic (Fryxell et al., 1981); Heard I. (Manguin, 1954); Weddell-Scotia
Confluence (Garrison et al., 1987); Fryxell et al. (1981) also described an Arctic form of this
bipolar species.

Large cells observed in the current study had only one row of marginal processes. The
smaller cells, which also resemble T. scotia (see below), have 2 rings of marginal strutted
processes with the labiate process formed between the rings. None of the cells observed
showed any evidence of occluded processes described for the typical T. antarctica and
T. scotia by Johansen & Fryxell (1985). The smaller cells are similar to T. scotia in the

formation of resting, heterovalvate cells. Hasle & Syvertsen (1997) and Fryxell et al. (1981)
outlined the reasons for the earlier segregation of T. antarctica var. borealis.

Thalassiosira australis Perag. (Peragallo, 1921: 84, pi. 4, fig. 17)

SYNONYMS: Coscinosira antarctica Mangin (1915: 55-57, fig. 39; pi. 1, fig. 8)

Actinocyclus excentricus Perag. (Peragallo, 1921 : 75-76, pi. 6, fig. 6)

Melosira dougetii Manguin (1960: 236, pi. 1, fig. 4)

ADDITIONAL REFERENCES: Fryxell (1977: 96-100, figs 1-12), Johansen & Fryxell (1985: 158, figs
19, 20, 46-48), Syvertsen (1985: 116-118, figs 15-27), Fryxell & Johansen (1990: 99, pi. 11.9,
figs 5, 6)

Fig. 2.45a-f

description: Cells usually by threads extruded from strutted

in short chains, closely linked
process; resting cells common Valves circular, 23-60 um diameter, markedly
in the chain.
convex (Fig. 2.45a, b). Areolation radial to fasciculate; areolae 1 1-13 in 10 um in the centre,
increasing to 14-16 in 10 um near the margin, loculate, opening externally via a simple
foramen, internally via an evenly pored velum (Fig. 2.45b). Strutted processes 4 or 5 on the
mantle, in clusters in the centre, often in 3 or 4 pairs of clusters, 5 in 10 um on the single
marginal ring. Marginal ring of strutted processes ranging from 7-9 in 10 um in resting spore
valves to 3 in 10 jam for rudimentary valves. External openings of resting spore strutted
processes long trumpet-shaped tubes (Fig. 2.45d), absent in rudimentary valves (Fig. 2.45f)-
Labiate process solitary, just inside marginal ring of strutted processes; internal opening
almost sessile and orientated along the radius. Occluded processes present on vegetative
valves (Fig. 2.45b) in a marginal ring inside the marginal ring of strutted processes, 2-3 in
10 um; external tubes of occluded processes ±straight, 2-7 um long. Girdle with areolate
valvocopula, a largely unstructured copula with or without a line of poroids adjacent to the

90
 DIATOMS

valvocopula, and 2-4 unstructured pleurae (Fig. 2.45f) with the ligulae rotating to the right.
For a complete description, see Fryxell (1977).

distribution: Inshore areas of Prydz Bay, near Davis Station, East Antarctica (e.g. Plough
I., 68°32'S 78°00'E and Ellis Fjord, 68°37'S 78°00'E); type locality, Petermann I., Antarctic

Peninsula, 65°09.8'S 66°32.7'W (Peragallo, 1921; not southern Argentina as suggested by

Fryxell, 1977); South Orkney Is. (Fryxell, 1977); Weddell Sea (Syvertsen, 1985); coastal
waters off Syowa Station, East Antarctica (Ishikawa et ah, 2001).

Slight differences were observed between material examined in this study and previous
descriptions. Strutted processes seen by us were often in 3 or 4 pairs of central clusters;
Syvertsen (1985) reported or 2 cluster pairs, and Fryxell (1977) reported 3 or 4 simple
1

clusters. Occluded processes present on vegetative valves in this study were 2-3 in 10 urn,
with external tubes straight and 2-7 urn long. Syvertsen (1985, figs 15, 21-23) illustrated
reflex-angled tubes 10-12 um long.

The resting spore is T. gerloffii (as mentioned by Syvertson, 1985:

similar to the valve of
119), but it differs from the having the labiate process in the same line as the
latter in not
marginal ring of strutted processes; in having trumpet-shaped, external tubes to the strutted
processes rather than irregularly linear ones; and a lower areolar density. The colony form
illustrated by Rivera (1981: figs 168, 179) for T. gerloffii shows cells attached by a central

Br m
M £

Figure 2.44. Thalassiosira antarctica. a, SEM External valve view of large, weakly silicified
form, b, SEM General internal view of large valve, c, SEM Internal view, detail of the
central cluster of strutted processes, d, SEM Internal view, detail of marginal strutted
processes and elongate labiate process. Scale bars: a, b = 5 um; c, d = 500 nm.

91
 DIATOMS

Figure 2.45. Thalassiosira australis. a, SEM

Vegetative valve, external valve view with
clusters of strutted processes (arrows) on the valve face, b, SEM Vegetative valve, internal
valve view of broken valve with the marginal ring of strutted processes (arrows) and
occluded processes, c, SEM Valves of adjacent cells still attached by threads, d, SEM
Resting spore, external valve view with the central cluster and marginal ring of strutted
processes and the single, longer external tube of the labiate process (arrow), e, SEM Oblique
view of cell with upper resting spore valve and lower vegetative valve showing vegetative
girdle structure, f, SEM Oblique view of cell with left-hand vegetative valve and right-hand
rudimentary valve showing similarity of girdle structure to that of the vegetative valve. Scale
bars: a, d-f = 5 urn; b = 1 urn; c = 10 urn.

92
 DIATOMS

Figure 2.46. Thalassiosira dichotomica. a, LM Valve, b, SEM Internal valve view showing
internal openings of marginal processes, single labiate process (arrow) and apparent lack of a
central process, c, SEM External view of valve without central strutted process, d, SEM
External view of valve with 1 central process and tangential thickening of areolae walls;
single labiate process (arrow), e, SEM External view of valve with 2 central process and
radial thickening of areolae walls, f, SEM Oblique external view showing girdle structure;
labiate process (arrow). Scale bars: a =1 \im\ b-f = 5 |xm.

93
 DIATOMS

Figure 2.47. Thalassiosira species, a, b, T. ferelineata. a, SEM Valve with single central
process, b, SEM Detail of finely pored velum of the areolae, strutted processes along the
margin, and the solitary labiate process (lower-right corner), c-f, T. frenguellii. c. SEM
External valve view showing coarse areolation and small strutted process in valve centre, d,
SEM Oblique view of cell showing girdle and small marginal processes (arrows), e, SEM
Oblique valve view with chitinous strand from central process, f, SEM External valve view
with chitinous strand from central process. Scale bars: a, c-f = 5 urn; b =
1 urn.

94
 DIATOMS

strand two or more times as long as the pervalvar dimension of the frustule compared to one
or less for the Weddell Sea material and no between-cell distance for the Prydz Bay material
of the current study.

Thalassiosira dichotomies (O.G.Kozlova) G.A.Fryxell & Hasle (1983: 54-57, figs 1-14)
BASIONYM: Porosira dichotomica O.G.Kozlova (1962: 8, fig. 2; fig. 3, no. 3)

ADDITIONAL REFERENCES: Kozlova (1967: 56, figs 2, 11, as Porosira dichotomica), Johansen &
Fryxell (1985: 161, figs 21-23, 33-35), Fryxell & Johansen (1990: 99, pi. 1.9, fig. 7-9), Hasle
1 &
Syvertsen(1997: 63)

Fig. 2.46a-f

description: Cells apparently solitary, with chitinous strands only observed from the
marginal strutted processes. Valves flat to slightly depressed (Fig. 2.46d, f), 12-26 urn
diameter; valve face weakly silicified, difficult to observe by LM (Fig. 2.46a). Areolae in a
distinct tangential (Fig. 2.46d) or radial (Fig. 2.46e) arrangement on valve face; radial
areolae 25-30 in 10 pm; one elongate areola between margin and ring of strutted processes
(Fig. 2.46f). Areolae with thickened walls, loculate, opening externally via a simple foramen
and internally by an evenly pored velum (e.g. marginal areolation of Fig. 2.46b). Single
marginal ring of strutted processes, 3-7 in 10 urn. Central processes 1-3 (-6) (Fig. 2.46b-e).
Strutted processes with short external extensions (Fig. 2.46d) supported by thickening from
adjacent areola walls; internal opening operculate, almost sessile and orientated along the
radius (Fig. 2.46b). Labiate process solitary, just inside marginal ring of strutted processes
and between 2 strutted processes (Fig. 2.46b, d, f). Girdle with an areolate valvocopula, a
largely unstructured and narrow copula, and 2 or 3 unstructured pleurae (Fig. 2.46f). For a
complete description, see Fryxell & Hasle (1983).

distribution: ADBEX (Southern Ocean, south of Australia); type locality, Indian Ocean
sector of the Antarctic (Kozlova, 1967); Pacific and Atlantic sectors of the Southern Ocean
(Fryxell & Hasle, 1983); probably with a circum-Antarctic distribution.

A distinctive feature of T. dichotomica is the concentric pattern of thickened areolae walls

formed on the valve face (Fig. 2.46e). Material examined in the current study has a density
of 25 areolae in 10 pm; a higher density (25-30 in 10 pm) was reported by Johansen &
Fryxell (1985). Our material also has 6- 7 strutted processes in 10 pm and a variable number
(?0-) 1-3 of central processes (Fig. 2.46b-e), whereas Johansen & Fryxell (1985) reported
3-7 strutted processes in 10 pm and 1-6 central processes.

Thalassiosira eccentrica (Ehrenb.) Cleve (1904: 216)

BASIONYM: Coscinodiscus eccentricus Ehrenb. (Ehrenberg, 1841: 146)
SYNONYM: Coscinodiscus labyrinthus Roper (1858: 21, pi. 3, fig. 2)

See VanLandingham (1968: 894; 1978: 3994) for further details of synonymy.

ADDITIONAL REFERENCES: Ehrenberg (1843: 371, pi. 1/3, fig. 20; pi. 3/7, fig. 5, as excentricus),
Hendey (1937: 242, as excentricus), Cupp (1943: 52, fig. 14), Fryxell & Hasle (1972: 300, figs 1-
18), Hasle & Syvertsen (1997: 62, pi. 6, tab. 8)

description: Cells chain-forming, discoid; pervalvar axis from one-sixth to longer than cell
diameter. Valves flat, circular, 15-110 pm diameter (40-140 [imfide Hendey, 1937); mantle

low and rounded. Areolation coarse but finer towards the margin, tangential or in 6-8 ill-
defined sectors of curved and somewhat parallel rows; areolae hexagonal, 5-11 in 10 pm;
central areolae usually larger than marginal ones. Strutted processes in 2 marginal rings;
processes 2-5 in 10 pm, each with a short external extension. One ring of short pointed
spines further from valve margin. Labiate process solitary, near the margin.

distribution: Ross Sea (Watanabe, 1982; Andreoli et al, 1995); Atlantic Ocean (Hustedt,
1958; Makarova, 1978); Prydz Bay, East Antarctica (Boden, 1985); Southern Ocean, south of
Australia (Wood, 1960).

95
 DIATOMS

The marginal ring of spines is a distinctive feature of this species. Hasle & Syvertsen (1997)
noted the distribution of T. eccentrica as "cosmopolitan, exclusive [of] polar regions".
Although we have not found this species, we include it because of the Antarctic records
mentioned above.

Thalassiosira ferelineata Hasle & G.A.Fryxell (1977: 26-28, figs 46-53)

ADDITIONAL REFERENCE: Herzig & Fryxell (1986: 14, tabs 1, 2)

Fig. 2.47a, b

description: Cells solitary. Valves flat, with a narrow mantle flaring to a broad marginal
rim, 20-43 um diameter. Areolae ±linear except where opposite the labiate process;
additional rows of small areolae at the valve margin result in the marginal row curving;
areolae ranging from 6-7 in 10 um at the valve centre to 16-18 in 10 um at the margin and
15-20 in 10 um on the valve mantle, with areolar diameter at the margin less than half that at
the centre. Areolae hexagonal, loculate, opening externally via a circular foramen, internally
via a fasciculately pored velum (Fig. 2.47b). Strutted processes in a single marginal ring, 3-4
in 10 um, with short internal extensions, operculate. Labiate process solitary, formed one
areola-width nearer valve face than strutted processes, with a short internal extension
orientated along the tangent. Single central process (visible by LM) causing displacement of
the surrounding areolae (Fig. 2.47a). For a complete description, see Hasle & Fryxell (1977).

distribution: MacquarieI. (54°58'S 155°26'E); type locality, "South Pacific Ocean (3°05'S

02°56'S 84°48'W-84°55'W)" (Hasle & Fryxell, 1977); tropical waters of northern Chile,
18°20'S 74°16'W (Rivera, 1981); warm-temperate waters of the Gulf Stream in the North
Atlantic (Herzig & Fryxell, 1986).

We have found only rare, isolated valves (but no intact cells), as was also the experience of
Hasle & Fryxell (1977). The occurrence of this species near Macquarie Island is the first
record for the sub- Antarctic and may indicate some mixing of subtropical and cold waters in
this region (see also T. symmetrica).

Thalassiosira frenguellii O.G.Kozlova (1964: 168, pi. 1, fig. 5)

ADDITIONAL REFERENCES: Kozlova (1967: 58-59, fig. 6), Fryxell & Hasle (1979b: 19-20, figs
25-30), Johansen & Fryxell (1985: 161, figs 6, 64), Fryxell & Johansen (1990: 99, pi. 11.8, fig. 2)

Fig. 2.47c-f

description: Cells chain-forming by means of a chitinous strand from the central strutted
process(es) (Fig. 2.47e,f), usually heavily silicified; inter-cell distance at least twice the

length of the pervalvar axis. Valves circular, 12-25 um diameter, those of smaller cells
slightly concave; marginal striae 15-18 in 10 eccentric; areolae 7-16 in 10
um. Areolation
um in the centre, increasing to 9-18 in 10 um
near the margin, loculate, opening externally
via a simple foramen and internally via a finely pored velum. Strutted processes 1 or 2 in the
valve centre, a single ring of processes at the margin (Fig. 2.47d, arrows), 1-3 in 10 um;
processes with internal extensions having tricolumnar supports. Labiate process solitary,
simple, with a tangential orientation to the internal opening. In small cells the single labiate
process is usually 3 or 4 areolae away from the central strutted process. For a complete
description, see Hasle & Fryxell (1979b).

distribution: Coastal waters off Davis Station, East Antarctica; type locality, Indian Ocean
sector of the Antarctic (Kozlova, 1964); south-western Atlantic (Johansen & Fryxell, 1985).

In specimens examined for the current study, the labiate process is 3 or 4 areolae removed
from the central strutted process. In contrast, Johansen & Fryxell (1985) reported a distance
of 4 or 5 areolae, while Fryxell & Hasle (1979) illustrated a distance of 7 or 8 areolae.

Areolation has been reported as irregular to linear (Fryxell & Hasle, 1979b) or irregular to
eccentric (Johansen & Fryxell, 1985). The densities of areolae in our material (7-10 in 10
um in the centre, 12-13 near the margin) varies slightly from those reported by others: 7-16
in 10 um in the centre (Johansen & Fryxell, 1985) and 9-13 (Fryxell & Hasle, 1979b) or 15-

96
 DIATOMS

18 (Johansen & Fryxell, 1985) near the margin. A lower density of marginal, strutted
processes (1-2 in 10 um) was reported by Fryxell & Hasle (1979b) and Johansen & Fryxell
(1985). Moreover, Johansen & Fryxell (1985) described the strutted processes having
internal extensions with tricolumnar supports, one of the features that distinguishes this
species from the closely related T. frenguelliopsis which has operculate strutted processes.
Marginal areolar density is typically greater (24-30 in 10 um) in T. frenguelliopsis.

Thalassiosira frenguelliopsis G. A. Fryxell & J.R.Johans., in Johansen & Fryxell (1985:

168, figs 6, 67, 68, 71,81)

ADDITIONAL REFERENCE: Fryxell & Johansen (1990: 99, pi. 1 1.8, Fig. 2)

Fig. 2.48a-d

description: Cells chain-forming, drum-shaped in girdle view. Valves circular, 12-34 um

diameter; marginal striae 24-30 in 10 um. Areolation irregularly linear or eccentric; areolae
7.5-16 in 10 um in the centre, increasing to 18 in 10 um near the margin, loculate, opening
externally via a simple pore with finger-like projections (Fig. 2.48b). Strutted process
solitary in valve centre, a single ring of processes at the margin, 1-2 in 10 um (Fig. 2.48c);
processes with internal extensions, with opercula covering 4 basal pores (Fig. 2.48d). Labiate
process solitary, located midway between centre and margin, internally slit-like (Fig. 2.48d).

« ' • »
* •
*fc
*
* % m "

Figure 2.48. Thalassiosira frenguelliopsis. a, SEM SEM Detail of

External valve view, b,
external view of areolae, central strutted process on and labiate process on right, c, SEM
left
Internal valve view with central and marginal processes, d, SEM Detail of internal surfaces
of areolae, central strutted process on left and labiate process on right. Scale bars: a, c = 5
um; b, d = um.
1

97
 DIATOMS

Figure 2.49. Thalassiosira species, a, b, T. gerloffii. a, SEM External valve view showing
coarse areolation. b, SEM Oblique view of cell with slightly concave valve, c, d, T. gracilis
var. expecta. c, LM General view of valve, d, TEM Internal valve view, showing 4 marginal
strutted processes, solitary off-centre strutted process and slit-like labiate process. Scale
bars: a, b = 10 um; c. d = 2 urn.

98
 DIATOMS

distribution: Southern Ocean, south of Australia; type locality, 50°06'S 11°46'E (south-
western Atlantic Ocean); sub-Antarctic (Fryxell & Johansen, 1990).

Thallasiosira frenguelliopsis is closely related to T. frenguellii, the latter having coarser

marginal striae (15-18 in 10 urn) and tricolumnar support of the internal extensions of the
strutted processes. Other species with similar morphology include T. gracilis var. gracilis,
T. gracilis var. expecta and T. perpusilla.

Thalassiosira gerloffii Rivera (1981: 73-80, figs 164-183)

ADDITIONAL REFERENCES: Johansen & Fryxell (1985: 168, figs 18, 44-45), Fryxell & Johansen
(1990, tab. 11.3, pi. 11.9, fig. 4), Hasle & Syvertsen (1997: 70)

Fig. 2.49a, b

description: Cells discoid, coarsely silicified; pervalvar axis c. 15 fim. Valves circular, 16-
77 urn diameter (23-50 [im fide Hasle & Syvertsen, 1997). Areolation fasciculate; areolae
13-18 in 10 urn in the centre, 12-20 in 10 p.m near the margin. Strutted processes 2-9 in the
valve centre, a single ring of processes at the margin, (5-) 6-9 (-11) in 10 um; processes
with external extensions visible by LM (Fig. 2.49b). Labiate process solitary, in the marginal
ring of strutted processes.

distribution: Southern Ocean, south of Australia; type locality, South Pacific, 51°12'S
74°05'W, Chilean coastal waters, southern Atlantic Ocean (Rivera, 1981); Antarctic sector of
south-western Atlantic Ocean (Johansen & Fryxell, 1985).

Thalassiosira gracilis (G.Karst.) Hust. (Hustedt, 1958: 109, pi. 3, figs 4-7) var. gracilis
BASIONYM: Coscinodiscus gracilis G.Karst. (Karsten, 1905: 78, pi. 3, fig. 4)

SYNONYM: Coscinodiscus minimus G.Karst. (Karsten, 1905: 78, pi. 4, fig. 8)

ADDITIONAL REFERENCES: Johansen & Fryxell (1985: 168, figs 8, 58, 59), Fryxell & Johansen
(1990: 99, pi. 1.8, fig. 4), Moisan &
1 Fryxell (1993: 293), Hasle & Syvertsen (1997: 82, pi. 12)

Fig. 2.50a-f

description: Cells solitary or chain-forming, discoid, heavily silicified, with a thin chitinous
strand from the central strutted process; pervalvar axis 3.5-9.5 um. Valves flat, circular, 5-28
p.m diameter. Areolation coarse; areolae 8-12 in 10 um in the centre, 16-20 in 10 um
towards the margin, loculate, opening externally via a simple foramen and internally via a
finely pored velum (Fig. 2.50d). Strutted process solitary in valve centre or off-centre, a
single ring of processes at the margin, 3-4 in 10 um; processes with internal extensions,
operculate. Labiate process solitary, located c. one-third of the distance from margin to valve
centre (Fig. 2.50b-d).

DISTRIBUTION: Antarctic sectors of the Atlantic, Indian and Pacific Oceans (Johansen &
Fryxell, 1985); Ross Sea (Watanabe, 1982; Andreoli et al, 1995); Lutzow-Holm Bay
(Tanimura et al., 1990); sub-Antarctic (Hustedt, 1958); subtropics (Fryxell & Hasle, 1979a);
KERFIX, 50°40'S 68°25'E, Southern Ocean (Kopczyhska et al., 1998).

Thalassiosira gracilis (G.Karst.) Hust. var. expecta (VanLand.) G. A. Fryxell & Hasle
(1979a: 384-386, figs 23-28)

BASIONYM: Thalassiosira expecta VanLand. (VanLandingham, 1978: 3995)

SYNONYM: Thalassiosira delicatula auct. non Ostenfeld (1908), sensu Hustedt (1958: 1 10, figs 8-10)

ADDITIONAL REFERENCES: Johansen & Fryxell (1985: 170, figs 8, 60-63), Hasle & Syvertsen
(1997: 83, pi. 12)

Fig. 2.49c, d

description: Cells solitary or chain-forming, discoid, heavily silicified, with a thin chitinous
strand from the central strutted process. Valves flat, circular, 7-26 um diameter. Areolation

99
 DIATOMS

radial, coarse; areolae 14-15 in 10 um in the centre, 16-20 in 10 um towards the margin,
loculate, opening externally via a simple foramen and internally via a finely pored velum.
Strutted process solitary in valve centre or off-centre, with a single ring of processes at the
margin, 3-4 in 10 um; processes with internal extensions, operculate. Labiate process
solitary, located c. one-third the distance from margin to valve centre.

distribution: Widely distributed in Antarctic waters (Johansen & Fryxell, 1985).

Only slight differences in valve diameter and areolar density separate var. expecta from the
type variety.

Thalassiosira gravida Cleve (1896: 12, pi. 2, figs 14-16)

SYNONYM: Thalassiosira tcherniai Manguin (1957: 1 15, figs 9, 51)

ADDITIONAL REFERENCES: Hendey (1937: 239, pi. 11, fig. 10), Hasle & Heimdal (1968: fig. 23),

Newell &Newell (1977: 170), Johansen & Fryxell (1985: 170, figs 27, 43), Hasle & Syvertsen
(1990a: 87, pi. 11.5, fig. 4), Moisan & Fryxell (1993: 293), Hasle & Syvertsen (1997: 68-70, pi.
8)

Fig. 2.51a-f

DESCRIPTION: Cells chain-forming, with chitinous strands from central processes connecting
the cells, sometimes in mucilaginous colonies, discoid, rectangular in girdle view; pervalvar
axis 6-15.8 urn (8-25 um fide Hendey, 1937). Valves circular, flat, 17-62 pm diameter.
Areolation radial; areolae 16-20 in 10 um, hexagonal, with finger-like projections
(Fig. 2.51c, d). Strutted processes numerous (c. 7) in valve centre and scattered over valve
face and mantle (Fig. 2.5 le) (5-10 in 10 um; Hasle & Syvertsen, 1997); processes
operculate, with external extensions. Labiate process solitary, large, marginal (Fig. 2.51b).

distribution: Coastal waters off Davis Station, East Antarctica and Drygalski Glacier, Ross
Sea; Antarctic sectors of the Indian and Pacific Oceans (Johansen & Fryxell, 1985);
Weddell-Scotia Confluence (Garrison et al., 1987).

Syvertsen (1977) noted that T. gravida is not known to form resting spores, unlike other
closely related species.

Thalassiosira lentiginosa (Janisch) G. A. Fryxell (1977: 100, figs 13, 14)

BASIONYM: Coscinodiscus lentiginosus Janisch (1878), in Schmidt et al. (1874-1959: pi. 58, fig. 11)

additional REFERENCES: Johansen & Fryxell (1985: 170, figs 7, 49, 50), Fryxell & Johansen
pi. 11.8, fig. 3), Moisan &
(1990: 99, Fryxell (1993: 293), Hasle & Syvertsen (1997: 79, pi. 10)

Fig. 2.52a-f

DESCRIPTION: Cells solitary, discoid, narrowly rectangular in girdle view; pervalvar axis 8-25
um. Valves flat, circular, 29-120 um diameter. Areolation usually fasciculate; areolae 7-9 in
10 um over valve face, c. 15 in 10 um at the margin (Fig. 2.52b), loculate, opening
externally via a foramen, internally via a cribrum (Fig. 2.52f). Strutted processes scattered
evenly over valve face, resembling small areolae (usually not visible by LM), 3-4 in 10 um,
lacking internal and external tubes. Labiate process solitary, marginal (Fig. 2.52a, e, f),
internal opening slit-like, with radial orientation. Chloroplasts small, numerous.

distribution: Coastal waters off Davis Station, East Antarctica; type locality, Table Bay,
near Cape Town, South Africa (Janisch, 1878); Indian, Atlantic and Pacific sectors of the
Antarctic (Johansen &
Fryxell, 1985); KERFIX, 50°40'S 68°25'E, Southern Ocean
(Kopczyhska et al., 1998); Ross Sea (Watanabe 1982); Weddell-Scotia Confluence (Garrison
et al., 1987); Southern Ocean, south of Australia (Wood, 1960); Prydz Bay, SIBEX (Boden,
1985).

00
 DIATOMS

Figure 2.50. Thalassiosira gracilis var. gracilis, a, LM General view of valve, b, SEM General
view of valve, c, SEM Internal valve view; note single submarginal labiate process, and
internal extensions of central and marginal strutted processes, d, SEM Internal view of valve;
note finely pored velum, e, SEM Oblique view of cell with clear view of marginal processes, f,
SEM Oblique view of cell with view of girdle bands. Scale bars: a = 10 um; b-f = 1 um.

101
 DIATOMS

Figure 2.51. Thalassiosira gravida, a, LM Valve view, b, SEM Valve, internal view with
single labiate process (arrow), c, SEM External view with detail of areolae and cribra. d,
SEM Olique view of cell highlighting strutted processes and mantle, e. SEM Cell with
chitinous threads from central strutted processes, f. SEM Girdle view with detail of
areolation of girdle bands. Scale bars: a-f = 1 pm.

102
 DIATOMS

Figure 2.52. Thalassiosira lentiginosa. a, SEM Valve with coarse striation and labiate
process (arrow), b, SEM Fasciculate areolation. c, SEM Oblique valve view showing
marginal labiate process (arrow) and copulae. d, SEM Detail of external view of labiate
process (arrow), e, SEM Internal valve view with labiate process (arrow), f, SEM Internal
view of round areolae with fine cribra (large arrows), 6 strutted processes with 2-3 adjacent
buttresses to the central pore (small arrows), and single slit-like labiate process. Scale bars:
a, b, e = 10 urn; c = 5 urn; d, f = 1 urn.

103
 DIATOMS

Figure 2.53. Thalassiosira maculata. a, LM

Valve face, b, LM Valve face, c, SEM Valve
face with distinct labiate process (arrow), d. SEM
Valve face with marginal ring of strutted
processes, e, SEM Internal valve view with marginal ring of strutted processes and1process
on valve face, f, SEM Internal valve view with tubular extensions of strutted processes,
single laterally expanded labiate process (small arrow), and chambered surface of
valvocopula (large arrows), g, SEM Oblique view of internal valve and girdle bands. Scale
bars: a, b, g = 10 um; c-f = 5 urn.

104
 DIATOMS

Figure 2.54. Thalassiosira maculata. a, LM Internal valve view, b, SEM Valve with 2
strutted processes (arrows), each located c. one-third of valve radius from centre, c, SEM
Internal valveview with 2 strutted processes, d, SEM Oblique internal valve view 2 strutted
processes, and chambered surface of valvocopula (arrows), e, SEM Valve with 2 strutted
processes (small arrows), located in line with the marginal labiate process (large arrow), f,
SEM Oblique view of e with 2 strutted processes (large arrows), and 1 row of large pores
(small arrows) on valvocopula adjacent to valve chambered surface. Scale bars: a = 10 |im;
b-f = 5 urn.

105
 DIATOMS

Figure 2.55. Thalassiosira maculata. a, SEM Internal valve view with 3 strutted processes
(arrows), b, SEM External valve view with 3 strutted processes (small arrows), and single
marginal labiate process (large arrow), c, SEM Oblique internal valve view with 3 strutted
processes (small arrows), and single marginal labiate process (large arrow), d, SEM Internal
valve view with 3 strutted processes, e, SEM External valve view with 3 strutted processes.
f, SEM Internal valve view with many scattered strutted processes. Scale bars: a-f = 5 urn.

106
 DIATOMS

Thalassiosira leptopus (Grunow) Hasle & G.A.Fryxell (1977: 20, figs 1-14)
BASIONYM: Coscinodiscus (lineatus var.?) leptopus Grunow, in van Heurck (1883, pi. 131, figs 5, 6)

SYNONYMS: Coscinodiscus leptopus Grunow, in Rattray (1890a: 476)

Coscinodiscus lineatus Ehrenb. (Ehrenberg, 1841: 146, pi. 3, fig. 4; 1854: pi. 22, fig. 6), Hendey (1937: 242),
Simonsen (1974: pi. 9. figs 3, 4)
Coscinodiscus leptopus var. discrepans Rattray (1890a: 476, pi. 2, fig. 3)

Coscinodiscus pseudolineatus Pant. (Pantocsek, 1886: 73, pi. 9, fig. 77)

Coscinodiscus praelineatus Jouse (1968: 15, pi. 2, fig. 1)

ADDITIONAL REFERENCES: Boden (1985: 28), Hasle & Syvertsen (1984: 145; 1997: 75, pi. 10)

description: Cells solitary, discoid. Valve face flat, circular, 26-165 urn diameter. Areolae
4-7 in 10 urn, arranged in tangential rows; central areola large; areolae on mantle small and
irregularly arranged. Strutted processes absent from valve face, but small processes (not
visible by LM) present in 2 or 3 marginal rings and 3-8 in 10 um. Labiate process solitary,
marginal, large; also a marginal ring of short occluded processes.

distribution: Type locality, Caltanisetta, Sicily (Ehrenberg, 1854); Prydz Bay, East
Antarctica, SIBEX (Boden, 1985); Ross Sea (Watanabe, 1982); Southern Ocean, south of
Australia (Wood, 1960).
While this species is known from widespread localities, including the Indian, Atlantic and
Pacific Oceans (Hasle & Fryxell, 1977), it is not commonly reported from Antarctic waters.

Thalassiosira maculata G.A.Fryxell & J.R.Johans., in Johansen & Fryxell (1985: 170,
figs 13, 69, 70, 72-74, 77-80)

SYNONYM: Coscinodiscus bullatus auct. non Janisch (1888), sensu Hustedt (1958: 1 12, figs 26-28)

additional reference: Fryxell & Johansen (1990: 99, pi. 1 1.8, fig. 9)

Figs 2.53a-g, 2.54a-f, 2.55a-f

description: Cells chain-forming, drum-shaped in girdle view. Valves discoid, 23-71 um

diameter. Areolation fasciculate; areolae 8-12 in 10 um in the valve centre, up to 14 in 10
um near the margin. Strutted processes in a marginal ring of 2-4 in 10 um and 1 -several
scattered over valve face (Figs 2.54b, 2.55b, f), operculate, with internal tubular extensions

(Fig. 2.53e-g). Labiate process solitary, marginal, slightly raised (Fig. 2.53c). Valvocopulae
with 1 row of large pores adjacent to valve (Fig. 2.54f). Copulae linking with valvocopulae
by chambers (Figs 2.53f, g, 2.54d) and interlocking ligulae.

distribution: Macquarie I. and Southern Ocean, south of Australia; type locality, Antarctic
sector of the Indian Ocean, 46°04.7'S 49°19.0'E, "Marion Dufresene" MD-08, Station 63
(Johansen & Fryxell, 1985); South Atlantic, 51°29'S 0°08'E (Hustedt, 1958).

Thalassiosira margaritae (Freng. & Orlando) O.G.Kozlova (1962: 8, pi. 2, fig. 4)

BASIONYM: Coscinodiscus margaritae Freng. & Orlando (Frenguelli & Orlando, 1958: 124, pi. 12, figs 6, 7)

ADDITIONAL REFERENCE: Jouse et al. (1962: 69, pi. 5, fig. 13)

description: Cells with circular valves, 24-27 um diameter. Areolation coarse, radial;
areolae 8-9 in 10 um. Central area of valve with small scattered granules partly dispersed
among nearby areolae. Valve with broad radially striated margin; striae 12-14 in 10 um.
distribution: Type locality, Scotia Bay, Laurie I., South Orkney Is., "Series 31 and 32"
(Frenguelli& Orlando, 1958); Indian Ocean sector of the Antarctic (Kozlova, 1962; Jouse et
al., 1962).

This uncommon species has been rarely reported in recent times.

107
 DIATOMS

Thalassiosira oestrupii (Ostenf.) Hasle (1972b: 544, figs 1-10)

BASIONYM: Coscinosira oestrupii Ostenf. (Ostenfeld, 1900: 52)
SYNONYM: Thalassiosira antiqua var. septata Proschk.-Lavr. (Proschkina-Lavrenko, 1955a: 48, pi. 1, figs 12-
14; pi. 2, figs 1, 2; 1955b: pi. 67, fig. 14)

ADDITIONAL REFERENCES: Fryxell & Hasle (1980: 805), Johansen & Fryxell (1985: 173, figs 9,
75, 76), Fryxell & Johansen (1990: 99, pi. 1.8, fig. 5), Hasle & Syvertsen (1997: 83, pi. 12)
1

Fig. 2.56a-d

description: Cells chain-forming, united by chitinous threads from the off-central strutted
process, discoid to rectangular in girdle view; pervalvar axis 0.5-2 times valve diameter.
Valves circular, 7-60 urn diameter, flat to slightly convex. Areolation eccentric, coarse;
areolae 5-10 in 10 um in the centre, 6-12 in 10 urn nearer the margin. Strutted processes
trifultate and operculate, with internal extensions (Fig. 2.56b); process in an off-centre
1

position and 1 marginal ring of processes 6-8 in 10 um (not visible by LM). Labiate process
solitary, not marginal, internally flattened and parallel to margin (Fig. 2.56b).

distribution: Coastal waters off Davis Station, East Antarctica; cosmopolitan (Fryxell &
Johansen, 1990); in broken sea-ice, Weddell Sea, Pacific and Atlantic sectors of the Southern
Ocean (Fryxell & Hasle, 1980); Weddell-Scotia Confluence (Garrison et al, 1987).

Figure 2.56. Thalassiosira oestrupii. a, LM General view of valve, b, SEM Internal view of
valve; note central and marginal strutted processes with internal extensions, and labiate
process inside of margin, c, SEM General view of valve, d, SEM Oblique view of cell. Scale
bars: a-d = 5 um.

108
 DIATOMS

Figure 2.57. Thalassiosira species, a, b, T. oliveriana. a, LM General view of valve, b, LM

General view of valve, c, d, T. scotia. c, LM Chain of germinating auxospores. d, LM
Auxospore. Scale bars: a, b = 5 pm; c = 50 pm; d = 10 pm.

109
 DIATOMS

Thalassiosira oliveriana (O'Meara) I.V.Makarova & V.A.Nikolajev (1983: 89, pi. 1,

figs 1-11; pi. 2, figs 1-11)

BASIONYM: Actinocyclus oliverianus O'Meara (1877: 58, tab. 1, fig. 7)

SYNONYMS: Podosira (?) oliveriana (O'Meara) Grunow, in van Heurck (1880-85: tab. 118, fig. 5)

Micropodiscus oliveriana (O'Meara) Grunow, in van Heurck (1880-85: tab. 1 18, fig. 5)

Actinocyclus antarcticus Castrac. (Castracane, 1886: 145)

Actinocyclus umbonatus Castrac. (Castracane, 1886: 145, tab. 4, fig. 4)

Schimperiella antarctica G.Karst. (Karsten, 1905: 88, tab. 8, fig. 6)

Schimperiella valdiviae G.Karst. (Karsten, 1905: 88, tab. 8, fig. 7a, b)

Actinocyclus antarcticus G.Karst. (Karsten, 1905: 91, tab. 9, fig. 2), nom. illeg.

Actinocyclus bifrons G.Karst. (Karsten, 1905: 92, tab. 9, fig. 8)

Actinocyclus janus G.Karst. (Karsten, 1905: 92, tab. 9, fig. 7)

Actinocyclus similis G.Karst. (Karsten, 1905: 92, tab. 9, fig. 4)

Actinocyclus valdiviae G.Karst. (Karsten, 1905: 92, tab. 9, fig. 3)

Actinocyclus excentricus Perag. (Peragallo, 1921: 75, tab. 6, fig. 6)

Schimperiella oliveriana (O'Meara) Sournia, in Sournia et al. (1979: 191, figs 8, 9), as oliverana

ADDITIONAL REFERENCES: Kopczyriska & Ligowski (1985: tab. 2), Priddle & Fryxell (1985: 146,
figs A, B), Fryxell & Johansen (1990: 99)

Fig. 2.57a, b

description: Cells discoid. Valves circular, 23-60 urn diameter. Areolation radial to
irregular, fine to coarse; areolae 7-8 in 10 um in the centre, 9-14 in 10 urn nearer the
margin; marginal band broad, distinctive, with fine areolation. Strutted processes in 1

marginal ring within marginal band, 3 in 10 um. Labiate process solitary, large, near the
margin.

distribution: Antarctic and sub- Antarctic (Fryxell & Johansen, 1990); Drake Passage
(Kopczyhska & Ligowski, 1985); Heard I. (Manguin, 1954); Cape Margerie, 66°50'S
141°25'E, Terre Adelie (Manguin, 1960, both as as Actinocyclus oliverianus).

The broad, marginal band of striae is a conspicuous feature of this species. Many early
Antarctic records were cited as Schimperiella oliverana, and the epithets oliverana and
oliveriana appear to have been used interchangeably.

Thalassiosira partheneia H.-J.Schrad. (Schrader, 1972: 59, pi. 1, 2)

ADDITIONAL REFERENCES: Hasle (1983: 223, figs 19-36), Hasle & Syvertsen (1997: 57, tab. 7)

Fig. 2.58a-d

description: Cells chain-forming, united by chitinous threads from the off-central strutted
process, sometimes in gelatinous masses, discoid to rectangular in girdle view; pervalvar axis
equal to or greater than valve diameter. Valves circular, 6-16 um diameter, flat to slightly
convex, thin-walled so that cells collapse easily, with radial ribs or rows of fine areolae (not
visible by LM). Areolae 40-60 in 10 um on the valve face and mantle. Areolation of copulae
and valvocopulae fine, irregular (Fig. 2.58c). Strutted processes in one marginal ring, 3-5 in
10 um, operculate, each with an external (Fig. 2.58b) and internal extension; 1 strutted
process in off-centre position, occasionally with 4 or 5 smaller satellite pores (Fig. 2.58d).
Labiate process solitary, midway between 2 of the marginal strutted processes (Fig. 2.58a,
b), internally slit-like and radially orientated.

distribution: Southern Ocean, south of Australia; type locality, upwelling area off Cape
Blanc, North Atlantic (Schrader, 1972); offshore West Africa and Peru (Hasle, 1983);
"mainly warm water region" (Hasle & Syvertsen, 1997); Gulf of Carpentaria and North-West
Shelf, Australia (Hallegraeff, 1984).

This is apparently the first record of this species from polar waters.

110
 DIATOMS

Figure 2.58. Thalassiosira partheneia. a, TEM Valve face with faint radial ribs, central and
marginal strutted processes, and marginal labiate process (arrow), b, SEM Valve face with
faint radial ribs visible near margin, central and marginal strutted processes, and marginal
labiate process (arrow), c, SEM Resting spore, d, TEM Off-centre strutted process (arrow)
with 4 smaller adjacent satellite pores. Scale bars: a-c = urn; d = 500 nm.
1

Thalassiosira perpusilla O.G.Kozlova (1962: 7, pi. 2, fig. 8)

ADDITIONAL REFERENCES: Fryxell & Hasle (1979a: 380, figs 1-11), Johansen & Fryxell (1985:
173, figs 5, 53-55), Fryxell & Johansen (1990: 99, pi. 1 1.8, fig. 1), Hasle & Syvertsen (1997: 83)

Fig. 2.59a-d

description: Cells lightly silicified, discoid in girdle view; pervalvar axis 0.5-1.5 times cell
diameter. Valves circular to slightly elliptical, 4-15 urn diameter, rounded towards the
margins. Areolae in irregular radial rows, 18-24 in 10 |xm; foramina partly occluded
(Fig. 2.59a). Areolation on valvocopulae finer than on valve face (Fig. 2.59c). Strutted
processes in a single marginal ring, 1-2 in 10 urn, operculate, each with a long internal
extension (Fig. 2.59b), lacking an external extension; strutted process in off-centre
1

position; strutted processes with 4 (rarely 3) satellite pores at their base (Fig. 2.59b). Labiate
process solitary, midway between 2 of the marginal strutted processes (Fig. 2.59a, b),
internally slit-like and radially orientated (Fig. 2.59b).

Ill
 DIATOMS

Figure 2.59. Thalassiosira perpusilla. a, SEM External view of valve face with off-centre
strutted process and eccentric labiate process (arrow), b, SEM Internal view of valve with 3
marginal and off-centre strutted process, and eccentric labiate process with flared opening
1

(arrow), c, SEM Girdle view of cell, d, SEM Girdle view of cell with highly developed
copulae. Scale bars: a-d = 1 urn.

distribution: Coastal waters near Davis Station, East Antarctica; type locality, Indian sector
of the Antarctic region between 20°E and 130°E, Second Expedition of the "Ob" (Kozlova,
1962); Antarctic and sub-Antarctic (Fryxell & Johansen, 1990); Weddell-Scotia Confluence
(Garrison et ai, 1987); Antarctic zones of the Weddell and Ross Seas and the Indian Ocean
(Fryxell & Hasle, 1979a).

Small cells (4-10 urn diameter) usually have only 2 or 3 marginal processes, but larger cells
(13-15 um diameter) can have 5 or 6 (Fryxell & Hasle, 1979a).

Thalassiosira poro-irregulata Hasle & Heimdal (1970: 573-575, figs 55-64, 71, 72)

ADDITIONAL REFERENCES: Makarova (1978: 224, figs 6-12), Rivera (1981: 113-116, figs 317-
339), Johansen & Fryxell (1985: 175, fig. 11), Fryxell & Johansen (1990: 99, pi. 11.8, fig. 7)

description: [after Johansen & Fryxell (1985)] Cells discoid. Valves circular, 21-30 urn
diameter (17-40.5 um fide Rivera, 1981), rounded towards the margins. Areolation
fasciculate; areolae 10-15 in 10 um. Marginal strutted processes in a single ring, 4-6 in 10
um. Central strutted processes 1-8, scattered in central area, each with an internal extension

12
 DIATOMS

with tricolumnar supports, lacking an external extension. Labiate process solitary, midway
between the valve centre and margin.

DISTRIBUTION: Antarctic and sub-Antarctic (Fryxell & Johansen, 1990); Antarctic, and
coastal waters of Chile and Peru (Johansen & Fryxell, 1985); Atlantic Ocean, 19°51.1'S
12°37'E, off Africa (Makarova, 1978).

We have not seen this species.

Thalassiosira poroseriata (Ramsfjell) Hasle (1972b: 544)

BASIONYM: Coscinosira poroseriata Ramsfjell (1959: 175, pi. 1, fig. g; pi. 2, fig. a)

ADDITIONAL REFERENCES: Hasle & Heimdal (1970: 574, figs 65-69), Rivera (1981: 117-120),
Johansen & Fryxell (1985: 175, figs 10, 51, 52), Fryxell & Johansen (1990: 99, pi. 1 1.8, fig. 6),

Hasle & Syvertsen (1997: 87, pi. 12)

Fig. 2.60a-f

description: Cells chain-forming, lightly silicified, discoid in girdle view; pervalvar axis
often longer than cell diameter. Valves 14-38 urn diameter, gently rounded
flat, circular,
towards the margins. Areolation radial or fasciculate; areolae 11-16 in 10 urn; areolation
finer on valve mantle (Fig. 2.60b). Marginal strutted processes in a single ring, or 2 in 1

10 urn. Central strutted processes 1-8, occurring in 1 or 2 straight or zig-zag lines, a central
areola or annulus at the end of the line (Fig. 2.60d, e); strutted processes with internal tube
extensions and tricolumnar supports. Labiate process solitary, not marginal, almost in line
with central strutted processes, internally slit-like (Fig. 2.60e, f).

distribution: Prydz Bay and Southern Ocean, south of Australia; cosmopolitan (Hasle &
Syvertsen, 1997); all sectors of the Antarctic (Johansen & Fryxell, 1985); sub-Arctic to
temperate northern waters (Fryxell & Johansen, 1990).

Thalassiosira ritscheri (Hust.) Hasle, in Hasle & Heimdal (1970: 569, figs 39-43)
BASIONYM: Coscinodiscus ritscheri Hust. (Hustedt, 1958: 1 17, figs 44-46)

ADDITIONAL REFERENCES: Johansen & Fryxell (1985: 176, figs 14, 56, 57), Fryxell & Johansen
(1990:99, pi. 11.8, fig. 10), Hasle & Syvertsen (1997: 86)

Fig. 2.61a-f

DESCRIPTION: Cells chain-forming by central threads, discoid in girdle view; pervalvar axis
c. 20 pm. Valves slightly convex, circular, 42-72 urn diameter. Areolation fasciculate, linear
or sublinear; areolae 12-16 in 10 pm, open to the outside by circular foramina with small
finger-like projections (Fig. 2.61d). Marginal strutted processes in a single ring, 3-4 in 10
pm. Central strutted processes 1-many in a cluster (Fig. 2.61b-e), with small internal
extensions. Labiate process solitary, near central strutted processes, internally slit-like (Fig.
2.61e).

DISTRIBUTION: Southern Ocean, south of Australia; Antarctic sectors of the Atlantic and
Pacific Oceans, Ross Sea, Weddell Sea (Johansen & Fryxell, 1985); Liitzow-Holm Bay
(Tanimura et al, 1990); Weddell-Scotia Confluence (Garrison et al, 1987).

Thalassiosira scotia G.A. Fryxell & Hoban, in Fryxell et al. (1979: 358-369, figs 2-18)

ADDITIONAL REFERENCES: Johansen & Fryxell (1985: 176, figs 25, 26, 40-42), Fryxell &
Johansen (1990: 99, pi. 11.9, figs 11, 12)

Fig. 2.57c, d

description: Cells discoid, heavily silicified. Valves circular, 15-56 pm diameter.

Areolation fasciculate, radial or bifurcate; areolae 9-16 in 10 pm. Marginal strutted
processes barely discernible by LM due to silicification, in 2 rings, 6-8 in 10 pm in outer
ring, 2-4 in 10 pm in inner ring. Central strutted processes 8-25, clustered, with external
extensions. Robust occluded processes also present in a marginal ring, 2-5 in 10 pm. Labiate

113
 DIATOMS

process solitary, located within occluded process ring. Resting spore valves heavily
silicified; areolae 5-8 in 10 urn; marginal striae present, sometimes vestigial (Johansen &
Fryxell, 1985).

DISTRIBUTION: Southern Ocean, south of Australia; south-western Atlantic and Scotia Sea
near South Georgia (Johansen & Fryxell, 1985).

Thalassiosira symmetrica G. A. Fryxell & Hasle (1972: 312, figs 37-46)

Fig. 2.62a-f

description: Cells discoid; pervalvar axis c. 18-20 urn. Valves flat, rounded at the margins,
circular, 30-88 um diameter. Areolation eccentric, finer on valve mantle than on valve face
(Fig. 2.62e); areolae (3-) 5-7 in 10 um in the centre, (5-) 6-8 in 10 um near the margin,
with internal cribra (Fig. 2.62b). Single central areola somewhat larger, surrounded by an
inconspicuous rosette of 7 areolae, in contrast to hexagonal pattern on remainder of the
valve. Strutted processes scattered over valve face, as well as in single marginal ring, 4-6 in
10 um, each with a short spine-like external extension (Fig. 2.62b, f), and a short internal
extension (Fig. 2.62d); strutted processes with 5 satellite pores at their base (Fig. 2.62d).
Two labiate processes on valve margin, c. 160-170° apart, internally slit-like (Fig. 2.62d),
externally with a very long extension (Fig. 2.62e, f)-

distribution: Prydz Bay, East Antarctica; type locality, 54°41'S 150°12'E, O.R.S. "Eltanin"
Cruise 38, Station 10a, Southern Ocean, south of Australia (Fryxell & Hasle, 1972); Pacific
and Indian Oceans and Gulf of Mexico (Fryxell & Hasle, 1972).

Thalassiosira trifulta G.A.Fryxell, in Fryxell & Hasle (1979b: 16-19, figs 1-24)

ADDITIONAL REFERENCES: Johansen & Fryxell (1985: 176, figs 12, 65, 66), Fryxell & Johansen
(1990:99, pi. 11.8, fig. 8), Moisan & Fryxell (1993: 293)

Fig. 2.63a-f

description: Cells discoid to drum-shaped; pervalvar axis 10-14 um. Valves flat to slightly
concave, circular, rounded at the margins, 16-58 um diameter. Areolation linear to sublinear,
finer on valve mantle than on valve face (Fig. 2.63d); areolae 5-6 in 10 um in the centre, 6-7
in 10 um near the margins, with internal pored vela (Fig. 2.63e, f). Strutted processes 1-4 in
valve centre and with a single marginal ring, 2-3 in 10 um, usually not visible by LM; each
process trifultate, with a long internal extension (Fig. 2.63f)- Labiate process solitary, not
marginal, internally slit-like and parallel to margin (Fig. 2.63f). Valvocopula with an
indistinct septum (Fig. 2.63d, e) and a complex triradiate ligula (Fig. 2.63e).

distribution: Coastal waters off Davis Station, East Antarctica; Antarctic sectors of the
Pacific and Atlantic Oceans, sub-Antarctic Indian Ocean, north-western Pacific Ocean
(Johansen & Fryxell, 1985).

Thalassiosira trifulta can be distinguished from poroseriata by having coarser valve

T.

1-16 in 10 um), and from

areolation (T. poroseriata has fasciculate areolation, with areolae 1

T. oestrupii by having noticeably more sparse, marginal strutted processes (T. oestrupii has

6-8 in 10 um and eccentric valve areolation).

Thalassiosira tumida (Janisch) Hasle, in Hasle et al. (1971 : 326, figs 1-43, 46)

BASIONYM: Coscinodiscus tumidus Janisch (1878), in Schmidt et al. (1874-1959: pi. 59, figs 38, 39)

SYNONYMS: Coscinodiscus tumidus \ar.fasciculatus Rattray (1890a: 475)

Coscinodiscus trigonus G.Karst. (Karsten, 1905: 84, pi. 5, fig. 10)

Coscinodiscus quinquemarcatus G.Karst. (Karsten, 1905: 85, pi. 7, figs 6, 6a)

Coscinodiscus inflatus G.Karst. (Karsten, 1905: 85, pi. 7, figs 7, 7a)

Coscinodiscus incurvus G.Karst. (Karsten, 1905: 85, 7, figs 8, 8a)

Coscinodiscus chunii G.Karst. (Karsten, 1905: 86, pi. 7, figs 10, 10a)

Coscinodiscus suktilis auct. non Ehrenberg (1841), sensu G.Karst. (Karsten, 1905: 86, pi. 7, figs 11, I la)

14
 DIATOMS

Figure 2.60. Thalassiosira poroseriata. a, LM Cell showing fasciculated areolation. b, SEM

Oblique view of cell showing finer areolation of valve mantle, c, SEM Valve with 2 central
strutted processes and labiate process (arrow), d, SEM Valve with 4 central strutted
processes and labiate process (arrow), e, SEM Internal view of valve with 4 central strutted
processes and labiate process (arrow); note internal extensions of marginal strutted
processes, f, SEM Internal view of valve with 2 central strutted processes and labiate process
(arrow). Scale bars: a-f = 5 um.

15
 DIATOMS

Figure 2.61. Thalassiosira ritscheri. a, LM Cell showing fasciculated areolation. b, SEM

Short chain of cells connected by central threads; misshapen cells are an artefact of SEM
preparation, c, SEMValve view of cell showing fasciculated areolation. d, SEM Central
cluster of strutted processes and labiate process (arrow), e. SEM Internal view of central
cluster of strutted processes and labiate process (arrow), f, SEM Internal view of valve;
labiate process (arrow). Scale bars: a-f = 5 urn.

116
 DIATOMS

Figure 2.62. Thalassiosira symmetrica, a, LM Valve face with eccentric areolation. b, SEM
Valve face with eccentric areolation; 2 marginal labiate processes clearly visible (arrows), c,
SEM Internal valve view showing positions of strutted processes and the 2 marginal labiate
processes (arrows), d, SEM Internal view of rounded areolae partly occluded with small
cribra, 5 strutted processes (small arrows) each with 5 minute satellite pores, and 1slit-like
labiate process (large arrow), e, SEM Girdle view of cell; external extension of labiate process
(arrows), f, SEM Oblique view of cell, showing spine-like marginal strutted processes and 2
labiate processes (arrows). Scale bars: a, b = 10 urn; c, e, f = 5 um; d = 1 um.

117
 DIATOMS

Figure 2.63. Thalassiosira trifulta. a. LM Valve view of solitary cell, b, SEM External valve
view with 2 central strutted processes (arrows), c, SEM External valve view with 2 central
strutted processes (arrows), d. SEM Oblique valve view showing girdle band structure, e.
SEM Oblique internal valve view showing marginal and off-centre strutted processs, single
1

labiate process (small arrow), and triradiate ligula (large arrow), f. SEM Internal valve view
of cell with 2 central strutted processes and a slit-like labiate process (arrow). Scale bars: a,
b = 10 urn; c-f = 5 urn.
 DIATOMS

Figure 2.64. Thalassiosira tumida. a, LM Valve view, b, SEM External valve view showing
concave surface, c, SEM Internal valve view showing slit-like internal endings of 6 marginal
labiate processes, d, SEM Oblique valve view showing girdle band structure, e, SEM Internal
and external views of labiate process, f, SEM Internal view showing velum of rounded
areolae, and 2 operculate strutted processes. Scale bars: a-d = 10 urn; e, f =
1 urn.
 DIATOMS

Figure 2.65. Achnanthes brevipes. a, SEM Chain of cells in girdle view, b, SEM Solitary
view, c, SEM Copulae (arrows) from parent cell surrounding upper valve with
cell in girdle
pseudoraphe. d, SEM Lower valve with raphe. Scale bars: a-d = 5 um.

20
 DIATOMS

Coscinodiscus quadrifarius Manguin (1960: 251, pi. 24, figs 284, 285)

For additional synonyms, see Hasle et al. (1971: 326).

ADDITIONAL REFERENCES: Rivera (1981: 127, figs 378-389), Johansen & Fryxell (1985: 176, figs
28-32), Kopczynska & Ligowski (1985: 74), Fryxell & Johansen (1990: 99, pi. 11.9, fig. 14),
Moisan & Fryxell (1993: 293), Hasle & Syvertsen (1997: 70)

Fig. 2.64a-f

DESCRIPTION: Cells usually solitary. Valves circular, 21-137 urn diameter, often concave in
the centre. Areolation linear to fasciculate; areolae 8-9 in 10 pm, loculate, opening
externally via a simple foramen and internally via an evenly pored velum. Strutted processes
generally scattered over valve face, apart from a ring of 6 or more near centre, positioned
between areolae in heavily silicified coarse valves or taking the place of areolae in lightly
silicified valves; marginal strutted processes in 2 or 3 rings, 5 processes in 10 pm, usually
not visible by LM. Strutted processes operculate, usually triradiate but with operculum 1

usually absent from processes near valve margin (Fig. 2.640- Marginal strutted processes
with a small granule or spine formed on upper (near valve face) rim of external opening.
Labiate processes 6 or 7, formed on margin, difficult to see by LM (especially in heavily
silicified valves), with slightly flared external openings (Fig. 2.64b, e), often compressed to
form an oval parallel to the direction of an internal slit that is slightly oblique to the
pervalvar axis. For a complete description, see Johansen & Fryxell (1985) or Hasle &
Syvertsen (1997).

DISTRIBUTION: Southern Ocean, 54-6 1°S, south of Australia; coastal waters off Syowa
Station, East Antarctica (Ishikawa et al., 2001); type locality, described as "Antarktik"
(Schmidt et al., 1874 1959); circumpolar within the Antarctic and sub-Antarctic (Hasle et
al, 1971); Ross Sea (Watanabe, 1982); Weddell-Scotia Confluence (Garrison et al., 1987).

The degree of silicification of the tumida appears to be sensitive to the

cell walls of T.

availability of silica, silicification being most evident

in cells found near the shore and near
convergence waters and at its lowest in cells in the intervening ocean or in places where they
divide rapidly in bloom conditions. Thalassiosira tumida is structurally similar to T. gravida,
T. rotula A.Meunier and T. antarctica, but it lacks the cluster of central strutted processes

seen in these taxa.

Order Pennales

Pennate diatoms have their valve structure, including the striae, arranged more or less in
relation to an axis rather than a central point or annulus, as in Centrales. In most species,
cells display bilateral symmetry in apical view.

Family Achnanthaceae

Achnanthes brevipes C.Agardh (1824: 1)

See VanLandingham (1967: 11) for synonymy.

ADDITIONAL REFERENCES: Smith (1853-56: 27, pi. 37, fig. 301), Boyer (1927: 232), Hendey
(1951: 41, pi. 16, figs 9, 10), Hustedt (1958: 143), Hendey (1964: 174, pi. 28, figs 7, 8), Andreoli
etal. (1995: 472, fig. 5)

Fig. 2.65a-d

description: Cells solitary or chain-forming, attached by valve surfaces, heterovalvate,

genuflexed in girdle view on the transapical axis; apical axis 29-90 um; transapical axis 30-
40 pm. Valves linear to elliptical; lower valve concave, with a raphe; upper valve convex,
with a slightly eccentric pseudoraphe. Transapical striae straight, uniseriate; poroid areolae
7-8 in 10 pm; poroids with complex cribra. On R-valve, the transverse area around the inner
raphe-ends is non-areolate (fascia) (Fig. 2.65d). Girdle structure composed of punctate

121
 DIATOMS

annular segments. Chloroplasts usually 4 irregular or elliptical bodies arranged in pairs

around the girdle (Hendey, 1964).

distribution: Plough I., near Davis Station, Prydz Bay, East Antarctica; Ross Sea
(Watanabe, 1982); Terra Nova Bay (Andreoli et al., 1995); Kerguelen Is. (Heiden & Kolbe,
1928); cosmopolitan, Great Britain (Hendey, 1964).

Material collected from coastal waters near Davis Station in the current study has smaller
29-61 urn) than those of previous reports.
cells (apical axis

Achnanthes vicentii Manguin (1957: 124, pi. 5, fig. 26a-e)

SYNONYM: Achnanthes vicentii naviculacae Manguin (1960: 308, pi. 14, figs 162a, b, 163a, b)

ADDITIONAL REFERENCES: Everitt & Thomas (1986: 5, fig. 3D, as A. vincentii), Chalmers et al.
(1996: 29, pi. 8, figs a, b)

Fig. 2.66a-d

description: Cells solitary, elliptical to lanceolate, heterovalvate; apical axis 6-21 urn;
transapical axis 3.2-5 urn. Valves flat; upper valve with a pseudoraphe (Fig. 2.66c); lower

valve with a raphe (Fig. 2.66a, b). Raphe central; terminal fissures curved in opposite
directions (Fig. 2.66a). Transapical striae radiating slightly, 9-13 in 10 um, triseriate,
containing poroids. Chloroplasts 2, lying on either side of the raphe.

distribution: Renwick Glacier, Amanda Bay, East Antarctica; type locality, Cape Margerie,
Terre Adelie (Manguin, 1957); epiphytic on marine macroalgae, near Davis Station, East
Antarctica (Everitt & Thomas, 1986).

Chalmers et al. (1996) noted that the species name has often been written as Acnanthes
vincentii. Specimens in the current study, from Prydz Bay, comprised comparatively small
cells, with slightly coarser striation (9 striae in 10 um) than described by Manguin (1957;
12-13 in 10 um).

Cocconeis
Cocconeis cells are solitary and heterovalvate. One valve (the P-valve) is typically convex
and bears a pseudoraphe; the other (the R-valve) is concave (or at least less convex than the
P-valve) and bears a central raphe-sternum. Valve structure is usually of broad, non-areolate,
transapical costae alternating with striae (intercostal areas) that contain uniseriate or
multiseriate poroids (areolae). The arrangement of striae and costae on the P-valve often
differs from that on the R-valve.

Cocconeis costata W.Greg, var. antarctica Manguin (1960: 304, pi. 14, figs 154, 155a-c)

ADDITIONAL REFERENCES: Andreoli et al. (1995: 472, fig. 10, SEM of C. costata)

Fig. 2.67a, b

description: Cells solitary, elliptical; apical axis 13.5-27 um; transapical axis 9-13 p.m.
Transapical costae slightly radiating, comparatively broad. Intercostal areas composed of
double rows of alternating areolae (Fig. 2.67a). Raphe-valve slightly concave; R-valve costae
12 in 10 um, with a submarginal non-areolate band creating a discontinuity of costae towards
the margins. Raphe a simple fissure with approximately parallel sides with small nodules at
valve centre. Pseudoraphe-valve with a narrow pseudoraphe; P-valve costae c. 5 in 10 um.

distribution: Coastal waters near Davis Station, East Antarctica; Cape Margerie, Terre
Adelie (Manguin, 1960).

Cells collected in the present study were 13.5-18 um long (apical axis).

122
 DIATOMS

Figure 2.66. Aclmanthes vicentii. a, SEM Raphe-valve. b, SEM Oblique view of raphe-
valve. c, SEM Pseudoraphe-valve. d, SEM Broken cell revealing external surface of raphe-

valve and internal surface of pseudoraphe-valve. Scale bars: a-d = 2 um.

23
 DIATOMS

Figure 2.67. Cocconeis cos fata var. antarctica, Entomoneis kjellmanii var. kjellmanii and
Manguinea fusiform is. a, b, C. costata var. antarctica. Pseudoraphe valves, c, E. kjellmanii
var. kjellmanii. SEM Colony of cells (from Everitt & Thomas, 1986). d, M. fusiform is. Cells
in apical and transapical view (from Manguin, 1960, as Tropidoneis fusiformis). Scale bars:
a, b = 2 p.m; c, d = 10 \xm.

24
 DIATOMS

Figure 2.68. Cocconeis fasciolata. a, LM Solitary cell, b, SEM Oblique view of P-valve
with pseudoraphe. c, SEM Valve with pseudoraphe. d, SEM Detail of striae and interstrial
costae. Scale bars: a-c = 10 u,m; d =
1 urn.

125
 DIATOMS

Figure 2.69. Cocconeis schuettii. a, SEM P-valve. with pseudoraphe. b, SEM Detail of P-
valve and slit-like, transapical striae, c SEM R-valve with poroid areolae; note non-areolate
areas (arrows) near valve centre and at raphe ends. d. SEM Detail of R-valve internal
surface. Scale bars: a. c = 10 urn; b, d =
1 urn.

126
 DIATOMS

Cocconeis fasciolata (Ehrenb.) N.E.Br. (Brown, 1920: 232)

BASIONYM: Rhaphoneis fasciolata Ehrenb. (Ehrenberg, 1844b: 204; 1854: pi. 35A/22, fig. 16)

SYNONYMS: Cocconeis pacifica Grunow (1867: 1 1, pi. 1, fig. 10)

Cocconeis imperatrix A. W.F.Schmidt (1894: pi. 189, figs 3, 10, 15)

Cocconeis costata var. pacifica (Grunow) Cleve (1895: 182)

Cocconeis pinnata var. plena Perag. (Peragallo, 1921: 53, pi. 2, fig. 4)

Cocconeis imperatrix f. plena (Perag.) Freng. (Frenguelli, 1943: 230, pi. 1, figs 3, 4)

ADDITIONAL REFERENCES: Peragallo (1921: 51, as C. costata var. pacifica), Andreoli el al. (1995:
472, fig. 9), Everitt & Thomas (1986: 8, fig. 3F)

Fig. 2.68a-d

description: Cells solitary, elliptical; apical axis 15-160 pm; transapical axis 25-75 pm.
Transapical striae radiating, composed of poroid areolae arranged in alternating alveolus-like
structures (Fig. 2.68d). Raphe-valve slightly concave; R-valve transapical striae 4-6 in 10
pm near the margin, 5-7 in 10 pm near the centre; interstrial costae broad, slightly raised.
Raphe a simple fissure of ±parallel sides with central endings (external) slightly dilated, but
internally constricted where they diverge towards valve margin. Pseudoraphe-valve convex,
with a narrow straight pseudoraphe (Fig. 2.68b); P-valve costae 3-4 in 10 pm near the
margin, 5-6 in 10 pm near the centre.

distribution: Ellis Fjord, Plough I. and coastal waters near Davis Station, East Antarctica

(Everitt & Thomas, 1986); Booth-Wandel I., Cape Tuxen, Petermann I. and Port Lockroy (all
Antarctic Peninsula) (Peragallo, 1921).

Cocconeis infirmata Manguin (1957: 123, fig. 24a-c)

SYNONYM: Cocconeis curiosa Hust. (Hustedt, 1958: 143, figs 96-99)

ADDITIONAL REFERENCE: Manguin (1960: 305, pi. 14, figs 156, 157)

description: Manguin (I960)] Cells solitary, elliptical; apical axis 16-17 pm;
[after
transapical axis 1-12 pm. Transapical striae radiating, curved towards the poles, uniseriate;
1

areolae poroid. Raphe-valve slightly concave; R-valve transapical striae 13 in 10 pm. Raphe
a simple straight fissure. Pseudoraphe-valve convex; P-valve transapical striae 11 in 10 pm,
the outermost areola of each stria noticeably larger.

distribution: Cape Margerie, Terre Adelie (Manguin, 1960).

This taxon has rarely been reported and was not found by us.

Cocconeis schuettii van Heurck (1909: 18, pi. 2, fig. 29)

SYNONYMS: Cocconeis pellucida sensu A.Mann (1907: 331)

Cocconeis gautieri van Heurck (1909: 17, pi. 2, fig. 30), Frenguelli & Orlando (1958: 82)
Cocconeis litigiosa van Heurck (1909: 18, pi. 2, fig. 28)

Cocconeis schuettii var. minor van Heurck (1909: 19, pi. 2, fig. 32)

Cocconeis schuettii var. litigiosa (van Heurck) Perag. (Peragallo, 1921: 54)

ADDITIONAL REFERENCES: Peragallo ( 192 1 : 53), Manguin (1960: 305)

Fig. 2.69a-d

description: Cells solitary, elliptical; apical axis 50-110 pm; transapical axis 30-70 pm.
Transapical striae curved and radiating, uniseriate; areolae poroid to slit-like. Raphe-valve
slightly concave; R-valve transapical striae 12 in 10 pm
near the margin, 18 in 10 pm near
the centre. Raphe a simple straight fissure, and outer ends, and
expanded slightly at the inner
connecting with an arrow-shaped non-areolate area at outer ends (Fig. 2.69c). Pseudoraphe-
valve convex near margin, but concave in central area around pseudoraphe. Areas
immediately around pseudoraphe and from c. midway from centre to margin are non-areolate

127
 DIATOMS

(Fig. 2.69a); P-valve transapical striae 24-25 in 10 urn near the margin (Fig. 2.69b), 20-21
in 10 um near the centre.

distribution: In sea-ice and epiphytic on Ballia, coastal waters near Davis Station, East
Antarctica; Petermann I., Antarctic Peninsula (Peragallo, 1921).

Cocconeis type A
Fig. 2.70a-d

DESCRIPTION: Cells solitary, elliptical; apical axis 87 urn; transapical axis 57 urn. Raphe-
valve not seen. Pseudoraphe-valve convex near the margin, concave in central area around
pseudoraphe. Pseudoraphe narrow, but expanded centrally and at both ends; P-valve
transapical striae fine (not visible by LM), discontinuous, curved and radiating, uniseriate;
areolae slit-like, 20 in 10 um (Fig. 2.70c). Areas immediately around pseudoraphe and from
c. midway from centre to margin are finely punctate (Fig. 2.70b, d).

distribution: Coastal waters of Inexpressible I., Ross Sea, Antarctica.

Insufficient material was available to determine the identity of this sample.

Family Cymbellaceae

Amphora
Most species of Amphora described from Antarctic waters have been based on early
collections. Examination of material (especially by electron microscopy) is required in order
to clarify many taxa, while Round et al. (1990) pointed out that the genus itself requires
revision. In the present study, only A. racovitzae was identified with certainty. Three other
species were also found, but they remain unidentified.

Van Heurck (1909) listed the following species collected on the "Belgica" Expedition:
A. racovitzae van Heurck, A. angusta var. angustissima van Heurck, A. arcta A. W.F.Schmidt,
A. cymbelloides Grunow, A. mexicana var. schmidtiana van Heurck, A. peragallorum van
Heurck, A. peragallorum var. robusta van Heurck and A. proteus var. oculata Perag. Hustedt
(1958) described A. antarctica (similar to A. cymbelloides) from 69°43'S 06°30'E, off
Dronning Maud Land.

Peragallo (1921) listed the following species collected during Charcot's "Porquoi Pas?"
Expedition of 1908-10: A. bongrainii Perag., A. gourdonii Perag., A. charcotii Perag.,
minor
A. petermannii Perag., A. proteus W.Greg., A. racovitzae van Heurck, A. racovitzae var.
van Heurck, A. senonquei Perag., A. virgata 0strup, A. virgata var. crassa Perag.,
A. peragallorum van Heurck, A. turgida W.Greg., A. lanceolata Cleve, A. lanceolata
var. aperta Perag., A. lanceolata var. robusta Perag. and A. liouvellei Perag.

Amphora racovitzae van Heurck (1909: 8, pi. 1, fig. 3)

SYNONYM: Amphora racovitzae var. minor Perag. (Peragallo, 1921: 61)

ADDITIONAL REFERENCE: Manguin (1960: 327, pi. 18, fig. 210)

Fig. 2.71a-e

description: Cells solitary, elliptical, with truncate ends in girdle view; apical axis 31-100
um; transapical axis 8.4-15 um. Valves asymmetrical about the apical plane; dorsal side
arcuate; ventral side concave, orientated such that the raphes lie in the same side of the cell.
Raphes eccentric, biarcuate; internal endings simple. Striae uniseriate, coarse; dorsal striae 7
in 10 um; ventral striae 6-7 in 10 um. Areolae slit-like. Girdle bands numerous, plain (Fig.
2.71c).

distribution: Burton Lake, sea-ice and coastal waters of Plough I., Kazak I., near Davis

28
 DIATOMS

Figure 2.70. Cocconeis type A. a, SEM Solitary cell, showing P-valve. b, SEM P-valve end
of cell; note slit-like areolae along submargin and central pseudoraphe. c, SEM Slit-like
areolae near valve margin, d, SEM Slit-like areolae near valve centre. Scale bars: a = 10 um;
b-d = 1 |am.

129
 DIATOMS

Figure 2.71. Amphora racovitzae. a, b, A. racovitzae var. racovitzae. a, SEM Oblique view
of ventral surface, b, SEM Ventral suface view, c-e, A. racovitzae var. minor, c. Ventral
surface with eccentric raphes along ventral margin of each valve, d, SEM Cell in valve view.
e, SEM Cell in dorsal view. Scale bars: a-e = 10 um.

30
 DIATOMS

Figure 2.72. Amphora, Amphora type A, SEM Girdle (dorsal) view of cell, with numerous
a,
areolated girdle bands, b, c, Amphora type B. b, SEM Dorsal view of cell, c, SEM Detail of

girdle bands between valve poles, d, Amphora type C, SEM Detail of girdle bands between
valve poles. Scale bars: a, b, d = 5 urn; c = 1 pm.

13
 DIATOMS

Figure 2.73. Amphiprora kufferathii. a. LM Chain of cells; note single and bilobed
chlorplasts. b,SEM Chain of cells in broad girdle view, c, SEM Chain of cells spiralling,
showing cells lying in broad girdle view and narrow girdle view, d, SEM Valve view; note
raphe supported on the edge of the keel (arrows), e, SEM Valve apex with detail of striation
on valve and keel. Scale bars: a, b, d = 10 urn; c = 50 urn; e = 2 um.

132
 DIATOMS

Station, East Antarctica; type locality, 64°25'S 62°02'W, Drake Passage (van Heurck, 1909);
Petermann I., Antarctic Peninsula (Peragallo, 1921); Cape Margerie, Terre Adelie (Manguin,
1960).

Cells found in Burton Lake sea-ice and coastal waters near Davis Station were small, with
the apical axis 31-53 urn, the transapical axis 8.4-10 um, width in girdle view 20-29 urn and
striae 10-11 um. They were very much in the size range of the Amphora racovitzae
in 10
var. minor material reported by Peragallo (1921).

Amphora type A
Fig. 2.72a

description: Cells solitary, elliptical, with truncate ends in girdle view; apical axis 25 |am;
cell width (not transapex) 9.2 um. Valves asymmetrical about the apical plane; dorsal side
arcuate. Striae uniseriate; dorsal striae 26 in 10 um. Areolae rounded. Girdle bands
numerous, perforated; areolae 26 in 10 um.

distribution: Coastal waters off Davis Station, East Antarctica.

The single cell examined by us is similar to an illustration in Round et al. (1990: 602, fig. m).

Amphora type B

Fig. 2.72b, c

description: Cells solitary, elliptical, with truncate ends in girdle view; apical axis 34 \im;
cell width (not transapex) 21.7 um. Valves asymmetrical about the apical plane; dorsal side
arcuate. Striae uniseriate; dorsal striae 25 in 10 um. Areolae slit-like. Girdle bands
numerous, striated; striae 38 in 10 um.

distribution: Coastal waters off Davis Station, East Antarctica.

Amphora type C
Fig. 2.72d

DESCRIPTION: Cells solitary, elliptical, with truncate ends in girdle view; apical axis 20-22
um; cell width (not transapex) 10 um. Valves asymmetrical about the apical plane; ventral
side concave; dorsal side presumably arcuate. Dorsal striae uniseriate, 13-14 in 10 um;
areolae loculate. Ventral striae biseriate, 13-14 in 10 um; areolae poroid. Girdle bands
numerous.

distribution: Coastal waters, Plough I., near Davis Station, East Antarctica.

The single cell found in the present study resembles an illustration in Round et al. (1990:
601, fig. 1).

Family Entomoneidaceae

Amphiprora kufferathii Manguin (1960: 320, pi. 16, fig. 188)

Fig. 2.73a-e

description: Cells large, solitary, twisted about the valvar axis, compressed and usually
lying in girdle view (where rectangular in shape); apical axis 52-79 um; transapical axis
c. 10 (xm. Valves elliptical to lanceolate; apices rounded, with a prominent wing-like keel

supporting the raphe (Fig. 2.73d). Valve striae 17-22 in 10 um (valve) and 20-24 in 10 um
(keel) (Manguin, 1960). Copulae numerous, open, porous. Chloroplast single, plate-like and
bi-lobed, or 2 per cell.

distribution: Prydz Bay, East Antarctica; Lutzow-Holm Bay (Tanimura et al., 1990);
Weddell-Scotia Confluence (Garrison et al., 1987).

133
 DIATOMS

Material collected for the current study has 17-22 striae in 10 urn (valve) and 22-24 striae in
10 urn (keel), with typically the same number of valve and keel striae on any given valve
face. This varies slightly from the 20 striae in 10 urn (valve and keel) originally described by
Manguin(1960).
Although Reimer & Patrick (1975) suggested that Amphiprora kufferathii should be
transferred to Entomoneis, no formal combination has been made.

Entomoneis kjellmanii (Cleve) Poulin & Cardinal (1983: 114, figs 23, 24)
var. kjellmanii

BASIONYM: Amphiprora kjellmanii Cleve, in Cleve & Grunow (1880: 15, pi. 4, fig. 83)
SYNONYM: Entomoneis kjellmanii var. typica Cleve-Euler (1952: 31), nom. inval.

See VanLandingham ( 1 967-79: 167) for details of synonymy.

ADDITIONAL REFERENCES: De Toni (1891: 334), Cleve (1894: 16), Meunier (1910: 291, pi. 32,
figs 4, 6, 7), Boyer (1927: 484), Hendey (1937: 349; 1964: 253), Everitt & Thomas (1986: 9, fig.

2F)

Fig. 2.67c

description: Cells large, solitary, twisted about the valvar axis, compressed and usually
lying in girdle view (where rectangular in shape); apical axis 60-130 urn; transapical axis
30-40 um. Valves elliptical to lanceolate; apices rounded, with a prominent wing-like keel
supporting the raphe. Valve striae 15-16 in 10 um (valve) and 1 1-12 in 10 um (keel) (Poulin
& Cardinal, 1983). Copulae numerous, open, porous. Chloroplast single, plate-like and bi-
lobed, or 2 per cell.

distribution: Type locality, Kara Sea, Arctic (Cleve & Grunow, 1880); Davis Station, Prydz
Bay, East Antarctica (Everitt & Thomas, 1986); Ross Sea (Andreoli et ai, 1995); Syowa
Station (Fryxell & Johansen, 1990); Weddell-Scotia Confluence (Garrison et ai, 1987).

Another variety, E. kjellmanii var. subtilissima van Heurck was recorded from 69°05'S
04°25'W and 69°43'S 06°30'E, Southern Ocean, Pacific sector (Hustedt, 1958).

Entomoneis paludosa W.Sm. var. hyperborea (Grunow) Poulin & Cardinal (1983: 1 16,
figs 29, 30)

BASIONYM: Amphiprora hyperborea Grunow, in Cleve & Grunow (1880: 62, pi. 5, fig. 86; 1894: 14)

SYNONYMS: Amphiprora paludosa f. minuta Grunow (1884: 106, pi. 1, fig. 51), Cleve-Euler (1952: 32,
fig. 1401c)
Amphiprora hyperborea var. genuina Cleve-Euler (1952: 32, fig. 1401a, b), nom. inval.

ADDITIONAL REFERENCE: McConville & Wetherbee (1983: 432)

Fig. 2.74a-d

description: Cells large, solitary, linear to lanceolate, twisted about the valvar axis,
compressed and usually lying in girdle view (where rectangular in shape); apical axis 29-123
um; transapical axis 12-16 um. Transapical striae 27-29 in 10 um on valve and keel.
Copulae numerous, open, porous.

distribution: Coastal waters off Davis Station and Polar Aviation I., East Antarctica; coastal
waters near Casey Station, East Antarctica (McConville & Wetherbee, 1983).

The type variety of E. paludosa has been recorded from the Ross Sea (Andreoli et ai, 1995,
as Amphiprora paludosa).

134
 DIATOMS

Figure 2.74. Entomoneis paludosa var. hyperborea. a, SEM Cell in girdle view, b, TEM
Detail of striae on keel and girdle bands, c, SEM Cell in girdle view, d, SEM Detail of striae
on valve. Scale bars: a, c = 10 um; b, d = 2 urn.

135
 DIATOMS

Figure 2.75. Fragilaria islandica. a-e, F. islandica var. islandica. a, SEM External valve
view, b, SEM External valve view, c, SEM Internal valve view. d. e. F. islandica var. stricta.
d, SEM Chain of cells, e, SEM Apical pore field on valve mantle. Scale bars: a-c = 5 (am; d =
10 \im; e = 2 urn.

136
 DIATOMS

Family Fragilariaceae

Fragilaria

Fragilaria is now regarded as an exclusively freshwater genus (see Williams & Round, 1987;
Round et al., 1990). Thus, the Antarctic species listed below (Fragilaria islandica and its

varieties) should probably be transferred to a different and possibly new genus. The apical
field structure of pores (cf. slits) helps to distinguish Fragilaria from Synedropsis.

Fragilaria islandica Grunow, in van Heurck (1880-85: pi. 45, fig. 37) var. islandica

ADDITIONAL REFERENCES: Lebour(1930: 193, fig. 152), Hendey(1964: 153)

Fig. 2.75a-c

description: Cells united to form flat ribbon-like colonies; cells and colonies rectangular in
girdle view; apical axis 20-50 pm; transapical axis 4-5 urn. Valves narrowly linear to
lanceolate, with subacute apices. Transapical striae fine, 22-24 in 10 pm, with a narrow
hyaline axial area that is more pronounced in the centre. Areolae poroid, uniseriate. Apical
pore field comprising short rows of areolae. Chloroplasts 2 narrow plates lying along the
valve.

distribution: Plough I. and Ellis Fjord, near Davis Station, East Antarctica; Arctic and
Norwegian coasts (Hendey, 1964).

Fragilaria islandica Grunow var. adeliae Manguin (1960: 298, pi. 12, fig. 128)

ADDITIONAL REFERENCE: Hasle et al. (1994: 255, figs 56, 76-78)

Fig. 2.76a-d

description: Cells united to form flat ribbon-like colonies; cells and colonies rectangular in
girdle view; apical axis c. 21-36 pm; transapical axis c. 3.5 pm. Valves narrowly linear to
lanceolate; valve ends elongate and subcapitate. Transapical striae 13-16 in 10 pm, with a
narrow hyaline axial area in the centre. Striae uniseriate; areolae poroid, c. 40 in 10 pm.
Apical pore field comprising short rows of areolae (Fig. 2.76e). Labiate process slit-like,
located at valve pole.

distribution: Ellis Fjord, near Davis Station, East Antarctica; Terra Nova Bay, Ross Sea;
type locality, Terre Adelie, 66°50'S 141°25'E (Manguin, 1960).

Cells found in the present study have valve areolae c. 40 in 10 pm and the apical pore field
of c. 5-8 x 3 pores on the valve mantle.

Material identified as F. islandica var. adeliae and reported for McMurdo Sound by Grossi
& Sullivan (1985) is possibly Synedropsis recta (see p. 144).

Fragilaria islandica Grunow var. producta Perag. (Peragallo, 1921: 68, pi. 3, fig. 7)

description: Cells united to form flat ribbon-like colonies; cells and colonies rectangular in
girdle view; apical axis 80-90 pm; transapical axis 4-5 pm. Valves narrowly linear to
lanceolate; valve ends elongate and capitate. Transapical striae 12 in 10 pm, with a narrow
hyaline axial area that is more pronounced in the centre. Striae uniseriate; areolae poroid.
Chloroplasts 2 narrow plates lying along the valve.

distribution: Type locality, Argentine Is., Antarctic Peninsula (Peragallo, 1921).

This variety has similar dimensions to Synedropsis recta. However, the pores in the apical
pore field of Synedropsis are slit-like, not poroid as in Fragilaria.

Fragilaria islandica Grunow var. stricta Perag. (Peragallo, 1921: 68, pi. 3, fig. 8)

Fig. 2.75d, e

137
 DIATOMS

description: Cells united by valve faces to form flat ribbon-like colonies; cells and colonies
rectangular in girdle view; apical axis 87-100 urn; transapical axis 6-7 um (3 urn at apices).
Valves narrowly linear to lanceolate. Transapical striae 14 in 10 um, with a narrow hyaline
axial area that is more pronounced in the centre. Striae uniseriate; areolae poroid. Areolae 50
in 10 um; pore field comprising c. 24
apical x 5 pores on the apical valve mantle.
Chloroplasts 2 narrow plates lying along the valve.

distribution: Inexpressible I., Ross Sea; type locality, Argentine Is., Antarctic Peninsula
(Peragallo, 1921).

Cells collected in the present study were 87 um long and c. 7 um wide in valve view.

Grammatophora arcuata Ehrenb. (Ehrenberg, 1853: 528)

See VanLandingham (1967-79: 1969) for details of synonymy.

ADDITIONAL REFERENCES: Grunow (1862: 420, tab. 11, fig. 7), Heiden & Kolbe (1928: 575)

Fig. 2.77a-d

DESCRIPTION: Cells solitary or in zig-zag colonies, arcuate in valve view; apical axis 22-39
um; transapical axis 6.3-8.8 um. Transapical striae straight, 15-16 in 10 um, each
comprising a uniseriate row of areolae extending down onto the mantle. Apical pore field
present at each valve pole (Fig. 2.77a). Two labiate processes per valve, 1 at each pole, lying
internal to the pore field (Fig. 2.77a, b). Fibulae 15-16 in 10 um (Fig. 2.77b). Copulae
numerous; valvocopula a complete ring (Fig. 2.77d).

distribution: Coastal waters off Davis Station, East Antarctica; Kerguelen Is. (Heiden &
Kolbe, 1928).

This species is usually found attached to macrophytes (e.g. Desmarestia in the present study)
or other substrata.

Pseudostaurosira brevistriata (Grunow) D.M.Williams & Round (1987: 276, figs 28-
31)
BASIONYM: Fragilaria brevistriata Grunow, in van Heurck (1880-85: 157, pi. 45, fig. 32)

Fig. 2.78a-d

description: Cells colonial, often united in short irregular chains by valve surfaces or poles,
rectangular in broad girdle view; apical axis 12-28 um; transapical axis 2-5 \im. Valves
slightly cruciform; surface with a broad somewhat ill-defined sternum and short spines along
the margin (8-19 in 10 um) which link adjacent cells; spines enlarged at the tips. Transapical
striae 13-17 in 10 um, uniseriate, with 2-4 areolae per stria at the centre, and fewer at the
poles; areolae comparatively large, circular to ovoid, sometimes extending below the spine
into the mantle. Apical pore fields small, consisting of a few pores. Copulae non-areolate,
numerous.

distribution: Plough I. and Ellis Fjord, near Davis Station, and coastal waters of Mawson
Station and Casey Station, East Antarctica; type locality, "Eaux douce - Bruxelles
(Delonge). Type no. 318" (Grunow, 1880).

Specimens collected by us closely resemble the Pseudostaurosira sp. of Williams & Round
(1987, fig. 36), being more elliptical than cruciate in valve outline. Cells from Plough Island,
Ellis Fjord and onshore Mawson coast were very variable in size and outline and were found
in considerable numbers during summer attached to seaweeds, mainly Chlorophyceae.

The smallest whereas the largest are more commonly

cells are usually circular, or nearly so,
it would be unwise
lanceolate. Intermediate forms produce such an uninterrupted series that
to assign them taxonomic rank. The assemblage of forms can only be regarded as a
polyphasic series.

138
 DIATOMS

Figure 2.76. Fragilaria islandica var. adeliae. a, SEM Oblique valve view, b, SEM End of
valve, c, SEM Chain of cells, d, SEM End of valve with apical pore field. Scale bars: a, c = 10

urn; b, d =1 urn.

39
 DIATOMS

Figure 2.77. Grammatophora arcuata. a, SEM Cell in valve view showing striae, 2
subapical labiate processes (small arrows) and apical pore fields (large arrows), b, SEM
Internal valve showing fibulae and the 2 subapical labiate processes, c, SEM Internal view of
valvocopula with undulate septa (arrows), d. SEM Half cell in oblique girdle and internal
valve view, showing septa and fibulae (in central valve area). Scale bars: a-d = 5 urn.

140
 DIATOMS

Figure 2.78. Pseudostaurosira brevisthata. a, SEM Solitary cell, apical axis, b, SEM Three
cells in valve view, c, SEM Two cells in transapical view, d, SEM Cell in transapical view
with short spines linking to adjacent cell (left). Scale bars: a-d = 1 urn.

41
 DIATOMS

Figure 2.79. Synedropsis species, a-c. S. hyperboreoides. a, SEM Stellate colony of cells
epiphytic on Entomoneis. b. SEM Cell in oblique valve view, c, SEM Apical field of minute
slits (arrow), d. e, S. laevis. d, SEM Single broken cell, e, SEM Apical field of slits, f, g. S.
recta, f, SEM Short chain of cells, g, SEM Internal valve view of apex with labiate process
(arrow). Scale bars: a. b, d, f = 5 uin; c. e. g = 1 urn.

142
 DIATOMS

Synedropsis

Some species formerly included in Fragilaria and Synedra have been transferred to
Synedropsis on the basis of their valve striation, the occurrence and structure of labiate
processes and apical pore (slit) fields and the nature of the girdle (Hasle et al., 1994).
Synedropsis hyperboreoides and S. laevis were collected in the present study, and several
other species, e.g. S. fragilis, S. recta, S. lata var. lata and S. lata var. angustata, have been
described from Antarctic locations (Hasle, 1994).

Synedropsis fragilis (Manguin) Hasle, Medlin & Syvertsen (1994: 265, figs 124-141,
142f, tab. 2)

BASIONYM: Synedra fragilis Manguin (1957: 123, pi. 5, fig. 22)

DESCRIPTION: [after Hasle et al. (1994)] Cells in stellate or pin-cushion-shaped colonies;

apical axis 26-58 urn; transapical axis 3-4 urn. Valves lanceolate, with obtuse ends; smaller
cellsrhomboidal. Transapical striae parallel to slightly radiating, 21-24 in 10 urn, visible by
LM. Areolae poroid, 50-60 in 10 um. Each valve end with an apical field of 6 or 7
longitudinal slits through the valve wall. Labiate process solitary, externally rectangular, c.
twice the size of an areola, located close to one valve end.

DISTRIBUTION: Type locality, Cape Margerie, Terre Adelie, 66°50'S 141°25'E (Hasle et al.,
1994). We have not seen this species.

Synedropsis hyperboreoides Hasle, Syvertsen & Medlin (1994: 251, figs 1-12, 17-21,
24-26,31-38,45-47, 142a)
ADDITIONAL REFERENCE: Hasle & Syvertsen (1997: 247)

Fig. 2.79a-c

DESCRIPTION: Cells in stellate colonies; apical axis 38-137 um; transapical axis 2.5-3 \im.
Valves linear to lanceolate with rostrate ends. Transapical striae parallel to slightly radiating,
35-40 in 10 um. Areolae poroid, 50-70 in 10 um. Each valve end with an apical field of 4-6
longitudinal slits through the valve wall. Labiate process solitary, close to one valve end.

distribution: Epiphytic on the large, sea-ice diatom Entomoneis, coastal areas of Davis
Station, East Antarctica; type locality, South Atlantic Ocean, 69°48'S 03°42'E, and coastal
Queen Maud Land and Weddell Sea (Hasle et al., 1994).

Synedropsis laevis (Heiden) Hasle, Syvertsen & Medlin (1994: 255, figs 61-67, 79-92,
142d)
BASIONYM: Cymatosira laevis Heiden, in Heiden & Kolbe (1928: 558, pi. 6, fig. 131)
SYNONYM: Nitzschia aciculariformis Manguin (1957: 132, pi. 7, fig. 45)

Fig. 2.79d, e

description: Only solitary cells observed. Valves lanceolate, tapering to rounded or rostrate
ends; apical axis 14-62 \im; transapical axis 3.5-5.6 um. Transapical striae parallel, 12-14
in 10 |xm. Areolae poroid. Each valve end with an apical field of 4-8 longitudinal slits

through the valve wall. Labiate process solitary, close to one valve end.

distribution: Coastal waters off Plough I., near Davis Station, East Antarctica; type
locality, Gaussberg, "Gauss" Expedition, 1902 (Heiden & Kolbe, 1928).

Synedropsis lata Hasle, Syvertsen & Medlin (1994: 261, figs 93-1 10, 142e, tab. 2) var.
lata

description: [after Hasle et al. (1994)] Cells solitary; apical axis 16-77 um; transapical axis
4-5.5 um. Valves narrowly elliptical; larger cells slightly dilated in central part. Transapical
striae parallel, 18-22 (occasionally 14-16) in 10 um, uniseriate; areolae poroid, 30-40 in 10
um. Sternum variable, narrow to broad. Apical fields of 5-8 longitudinal slits through the

143
 DIATOMS

valve wall at one end and 4-7 slits at the other, asymmetrically located on the valve. Labiate
process solitary, close to the latter valve end.

distribution: Type locality, Antarctic sea-ice, 60°42'S 46°36'W [IMBB Sample 58(12)]
(Hasle era/., 1994).

This diatom was not found in the present study. Another Antarctic variety, S. lata var.
angustata Hasle, Syvertsen & Medlin, has a more elongate valve outline, although the size
ranges overlap (Hasle et al., 1994). Var. angustata also has fewer apical field slits at the
labiate process end. Synedropsis lata is distinctive in having a narrowly elliptical to linear
valve outline, as well as asymmetrical apical fields with an unequal number of slits at each
apex.

Synedropsis recta Hasle, Syvertsen & Medlin (1994: 252, figs 27-30, 51-55, 57-60,
68-75, 142c)

Fig. 2.79f, g

description: [after Hasle et al. (1994)] Cells united at poles to form stellate or bundle-
shaped colonies. Valves linear to lanceolate, with rostrate or subcapitate ends; apical axis
17—48 um; transapical axis 3.5-5 urn. In girdle view, cells narrowly linear with deep valve
mantles. Transapical striae parallel, 11-14 in 10 um, uniseriate; areolae poroid. Sternum
broad. Each valve end with an apical field of 5-8 longitudinal slits through the valve wall.
Labiate process solitary, close to one valve end.

distribution: Coastal waters off Casey Station, East Antarctica; type locality, Antarctic sea-
ice, 75°35'S 26°46'W [IMBB Sample 2486] (Hasle et al, 1994); South Atlantic, Weddell
Sea, Terre Adelie, McMurdo Sound (see Hasle et al, 1994).

Hasle et al. (1994) included Fragilaria (?) "sp. a" Tanimura {in Tanimura et al., 1990: 28,
figs 4, 5, 7) as a synonym of S. recta.

Thalassionema nitzschioides (Grunow) Mereschk. (Mereschkowsky, 1902: 78)

BASIONYM: Synedra nitzschioides Grunow (1862: 403, pi. 5, fig. 18)

SYNONYMS: Thalassiothrix nitzschioides (Grunow) Grunow, in van Heurck (1881: pi. 37, figs 11-13; pi. 43,
figs 7-10)
Synedra nitzschioides var. minor Cleve (1883: 482)
Thalassiothrix curvata Castrac. (Castracane, 1886: 55, pi. 24, fig. 6)

Thalassiothrix frauenfeldii var. nitzschioides (Grunow) Jorg. (Jorgensen, 1900: 21)

Thalassionema nitzschioides f. curta [?error of F.W.Mills (1935: 1605) for Thalassiothrix curvata Castrac]

See VanLandingham (1978: 3991) for full synonymy.

ADDITIONAL REFERENCES: Heiden & Kolbe(1928: 563, pi. 5, figs 115-118), Hendey (1937: 336),
Hustedt (1958: 139), Hallegraeff (1986: 60-65), Hasle & Syvertsen (1997: 257, pi. 56(2), 57a, b,
tab. 66)

Fig. 2.80a-d

description: Cells straight, forming stellate or zig-zag colonies. Valves linear, tapering,
rounded towards the apices; apical axis 10-80 (-105) um; antapical axis 2-3.5 um; pervalvar
axis c. 2 (-7) um. Valve surface flat. Areolae located along margins, externally with
elaborate vela (Fig. 2.80b), internally simple, 10-12 in 10 um.

distribution: Southern Ocean, 60°50'S 93°36'E, offshore Wilhelm Land, coastal waters II

off Davis Station, East Antarctica, and southern Pacific Ocean; Kerguelen Is. (Heiden &

Kolbe, 1928); coastal Europe and North Atlantic Ocean (Hendey, 1937); "cosmopolitan but
not in the high Arctic and Antarctic" (Hasle & Syvertsen, 1997).

One unusually large cell in the present study measured 105 * 7 um, but otherwise it had all

the attributes of T. nitzschioides. A similar species, T. gelida Perag., described from

Petermann Island, has cells 100-120 um long and 8 areolae in 10 um.

144
 DIATOMS

Figure 2.80. Thalassionema nitzschioides. a, SEM Cell in valve view, b, SEM Elaborate
areolae on margins of valve surface, c, SEM Cell internal valve view, d, SEM Cell in girdle
view. Scale bars: a, c, d = 10 ^m; b = 1 \\.m.

45
 DIATOMS

Figure 2.81. Thalassiothrix antarctica. a, LM Single cell, b, SEM Single broken cell. c.
SEM Detail of marginal spines on frustule. d, SEM Head pole of cell with 2 protruding
spines, e, SEM Internal view of head pole of cell; note labiate process and marginal areolae.
Scale bars: a-c = 10 urn; d, e =1urn.

146
 DIATOMS

Thalassiothrix antarctica A.Schimp. ex G.Karst. (Karsten, 1905: 124, pi. 17, fig. 12)

SYNONYMS: Synedra thalassiothrix Cleve ex O.Miill. (Miiller, 1877a: 20)

Synedra tabulate var. thalassiothrix Schwarz ex O.Miill. (1877b: 165)

Thalassiosira antarctica var. echinata G.Karst. (Karsten, 1907: 396, pi. 46, fig. lOa-c)
Thalassiothrix longissima var. antarctica Cleve & Moller (1878, No. 125), nom. nud.
Thalassiothrix longissima var. antarctica Cleve & Grunow, in van Heurck (1881, pi. 37, fig. 10), HallegraetT

(1986: 64, figs 15-22)

ADDITIONAL REFERENCES: Hendey (1937: 335), Hustedt (1958: 141), Semina (1981: 147),
Hallegraeff(1986: 64, figs 15-22), Hasle & Semina (1987: 181, figs 26-59), Hasle(1990: 133, pi.
17.1, figs 4-6), Hasle & Syvertsen (1997: 266, pi. 58, tab. 67)

Fig. 2.81a-e

DESCRIPTION: Cells needle-like, straight, slightly curved or sigmoidal in girdle view, forming
radiating colonies, joined by bent foot poles (one end of frustule); head poles separated.
Apical axis 420-5680 urn; transapical axis 1.5-6 urn; foot pole apex rounded; head pole
apex rounded but with 2 protruding spines. Areolae in a single marginal row, 12-17 in 10
jam, with complex vela. Row of fine spines, marginal on the vela, pointed towards the head
pole, 1-2 in 10 um, occurring along the frustule but more abundant towards the head pole.
Small foramina on valve surface. Labiate process at each apex (see Hasle & Semina, 1987).
Chloroplasts numerous, spherical.

distribution: Heard I.; type locality, Antarctic Ocean, "Challenger" Expedition; Kerguelen
(Hustedt, 1958); KERFIX, 50°40'S 68°25'E, Southern Ocean (Kopczyhska et al, 1998);
Southern cold-water region (Hasle & Syvertsen, 1997); wide distribution in South Atlantic,
Pacific and Indian Oceans (Hendey, 1937).

Thalassiothrix antarctica can form extensive mats of intertwined cells which have been
mistaken for shallow rocks on marine charts (Quilty et al., 1985). Hasle (1990) noted that it
often occurs in association with Trichotoxon reinboldii. The closely related T. longissima
Cleve & Grunow is a northern, cold-water species.

Trichotoxon reinboldii (van Heurck) F.Reid & Round (1987: 224)

BASIONYM: Synedra reinboldii van Heurck (1909: 23, pi. 3, fig. 35)

SYNONYM: Synedra pelagica Hendey (1937: 335)

ADDITIONAL REFERENCES: Hasle & Semina (1987: 189, figs 67-75), Hasle (1990: 133, pi. 17.1,
Moisan & Fryxell (1993: 293), Hasle & Syvertsen (1997: 267)
figs 7-9),

Fig. 2.82a, b

DESCRIPTION: Cells needle-like, arcuate, tapering gradually towards the poles, solitary or in
colonies; apical axis 800-3600 um; transapical axis 3-10 urn; poles similar and without
spines (cf. the spine-bearing Thalassiothrix antarctica); striae on mantle, extending slightly
onto the valve face, 8-10 in 10 um; areolae with external cribra scattered over valve face.
Labiate process at each pole. Chloroplasts numerous, discoid.

distribution: Endemic to the Southern Ocean, possibly circumpolar (Hasle, 1968; Reid &
Round, 1987); Weddell Sea (Moisan & Fryxell, 1993); Indian Ocean sector of the Antarctic
(Kopczyhska et al., 1986); Weddell-Scotia Confluence (Garrison et al., 1987).

Hasle (1990) noted that this species often occurs in association with Thalassiothrix
antarctica (above): "They are approximately of the same length and width and are similar in
cell shape, with Trichotoxon reinboldii being curved, but not sigmoidal. Trichotoxon
reinboldii also forms colonies but, unlike Thalassiothrix antarctica, the cells are attached at
either end resulting in pointed, ovoid colonies (Reid & Round 1987)".

147
 DIATOMS

Family Naviculaceae

Banquisia belgicae (van Heurck) Paddock (1988: 79, pi. 28)

BASIONYM: Amphiprora belgicae van Heurck (1909: 14, pi. l.figs 11, 15)

SYNONYM: Tropidoneis belgicae (van Heurck) Heiden, in Heiden & Kolbe (1928: 655, tab. 4, figs 98, 99)

ADDITIONAL REFERENCES: Paddock ( 1990: 152, pi. 19.1, figs 5, 6), Hasle & Syvertsen (1997: 286,
pi. 64, tab. 72)

Fig. 2.82c, d

description: Cells usually united in ribbons, lanceolate in valve view; apical axis 70-125
um; pervalvar axis 21-37 um. Valves biarcuate in girdle view, with a depression at the
central nodule. Raphe on a raised flat valve surface, dividing the valve into 2 equal sides.
Raphe fins present. Transapical striae 16-20 in 10 um, punctate.
distribution: Coastal waters near Davis Station, East Antarctica; Weddell-Scotia
Confluence (Garrison et al., 1987); Liitzow-Holm Bay (Tanimura et al, 1990); in pack-ice
off Wilhelm II Land (Heiden & Kolbe, 1928).

Berkeleya rutilans (Trentep. ex Roth) Grunow (1880: 1587)

BASIONYM: Conferva rutilans Trentep. ex Roth (1806: 179)
SYNONYM: Amphipleura rutilans (Trentep. ex Roth) Cleve (1894: 126)

See VanLandingham (1967: 507) for further references and synonymy.

ADDITIONAL REFERENCES: Cardinal et al. (1984: 371, figs 9, 10, 13, 14), Lobban (1986: 85, fig.
1 1 ), Round et al. ( 1 990: 5 1 8, figs f, g)

Fig. 2.83d

description: Cells solitary or forming mucilage tubes. Valves linear to lanceolate, with
bluntly rounded poles; apical axis 13-60 um; transapical axis c. 9-12 um. Raphe slits short,
with central raphe endings terminating well short of valve centre; all raphe endings deflected
in the same direction. Transapical striae fine, straight, uniseriate, c. 30 in 10 um, consisting
of simple poroids. Copulae bearing 2 rows of round to oval poroids.

distribution: Ellis Fjord, near Davis Station, East Antarctica; high intertidal, Livingston I.,

Antarctic Peninsula (Lobban, 1986).

Specimens seen in the present study are longer (50-60 um) than those of previous reports
(e.g. 13-43 um, Cardinal et al., 1984). Amphipleura rutilans (Trentep. ex Roth) Cleve var.
antarctica (Harv.) Grunow, with finer valve striation, was reported by Manguin (1960: 309,
fig. 164) from the Antarctic Peninsula. " Berkeley alAmphipleura" was recorded by
McConville & Wetherbee (1983) from near Casey Station, East Antarctica.

Ephemera planamembranacea (Hendey) Paddock (1988: 86, pi. 31)

BASIONYM: Navicula planamembranacea Hendey (1964: 188, fig. 8)

additional REFERENCE: Hasle & Syvertsen (1997: 284, pi. 64)

Fig. 2.84a-f

description: Cells usually solitary, flattened transapically, usually seen in girdle view,
weakly silicified and often collapsed; apical axis 66-90 um; transapical axis 8-10 um;
pervalvar axis 15-30 um. Transapical striae fine, 28-30 in 10 um; striae near the centre of
cell visible by LM (Fig. 2.84b). Raphe located on a raised flat valve surface, dividing the
valve into 2 unequal sides. Central nodule slightly depressed (Fig. 2.84a, b). Raphe fins small,
located near valve ends on either side of the nodule. Chloroplasts small, spherical, numerous.

distribution: Southern Ocean, south of Australia; North Atlantic Ocean (Hasle &
Syvertsen, 1997).

48
 DIATOMS

Figure 2.82. Trichotoxon and Banquisia species, a, b, T. reinboldii. a, LM Central part of

cell; note smooth margins, b, LM Valve pole showing irregular pattern of areolae close to the
apex, c, d, B. belgicae, LM Solitary cells in girdle view. Scale bars: a-d = 10 um.

149
 DIATOMS

Figure 2.83. Haslea, Berkeleya and Membraneis species, a-c, H. trompii. a, SEM Solitary
cell invalve view, b, SEM Detail of central raphe endings (arrows), c, Navicula [Haslea]
trompii var. major, SEM Detail of cell apex, d, B. rutilans, SEM Solitary cell. e.
M. challengeri, LM Solitary cell. Scale bars: a, d. e = 10 u,m; b. c = 5 urn.

50
 DIATOMS

Figure 2.84. Ephemera planamembranacea. a, LM Cell in girdle view, b, LM Cell in girdle

view, c, SEM Cell in broad girdle view, d, SEM Internal view of valve pole with
helictoglossa. e, SEM Cell in broad girdle view, f, TEM Cell in broad girdle view. Scale
bars: a-c, e, f = 10 )a.m; d = 1 |j.m.

51
 DIATOMS

Figure 2.85. Gyrosigma sp. a, SEM Solitary cell in valve view, b, SEM Solitary cell in
oblique valve view, c, SEM Solitary cell in valve view, d, SEM Pole raphe ending, e, SEM
Central raphe endings. Scale bars: a-c = 20 urn: d. e = 1 urn.

52
 DIATOMS

This appears to be the first record of this species from southern waters.

Gyrosigma sp.

Fig. 2.85a-e

description: Cells solitary, usually lying in valve view. Valves linear to lanceolate,
sigmoidal; apical axis 170-210 um; transapical axis 25-50 urn; pervalvar axis, 20-40 urn.
Raphe sigmoidal, central on valve face, with terminal raphe endings turned in the same
direction as the valve pole; central raphe endings turned in the opposite direction. Transverse
and longitudinal striae fine; transverse striae slit-like, 26-28 in 10 urn; longitudinal striae
14-16 in 10 (am. Chloroplasts 2, plate-like.

distribution: Franklin I., Ross Sea.

Insufficient material was examined for an unambiguous identification. Species of Gyrosigma

have a similar raphe and valve system to those of Pleurosigma.

Haslea trompii (Cleve) Simonsen (1974: 47)

BASIONYM: Navicula trompii Cleve (1901a: 932, fig. 1 1)

ADDITIONAL REFERENCES: Heiden & Kolbe (1928: 625, pi. 3, fig. 77), Hasle & Syvertsen (1997:
278, pi. 62)

Fig. 2.83a, b

description: Cells solitary, weakly silicified. Valves lanceolate in valve view, tapering
towards the ends; apical axis 70-160 um; transapical axis 10-14 urn. Raphe central, straight;
central raphe pores simple, very close together; terminal raphe endings hook-like. Transverse
and longitudinal striae fine (but visible by LM), crossing at right angles; transverse striae
28-30 in 10 um; longitudinal striae 17-20 in 10 um. Chloroplasts 2, plate-like.
DISTRIBUTION: In pack-ice, c. 65°S 86°E, off Wilhelm II Land, "Gauss" Expedition (Heiden,
& Kolbe, 1928); Lutzow-Holm Bay (Tanimura et al., 1990); southern cold-water region
(Hasle & Syvertsen, 1997).

Navicula [Haslea] trompii var. major (Heiden & Kolbe, 1928) was also found in the present
study, at Terra Nova Bay (Ross Sea). This variety has the following dimensions: apical axis
258-378 um; transapical axis 20-27 um; transapical striae 20 in 10 um; and longitudinal
striae 11-12 in 10 um (Fig. 2.83c). Both varieties were reported from Antarctic pack-ice off
Wilhelm II Land by Heiden & Kolbe (1928).

Luticola muticopsis (van Heurck) D.G.Mann, in Round et al. (1990: 671)

BASIONYM: Navicula muticopsis van Heurck (1909: 12, pi. 2, fig. 181)

SYNONYMS: Navicula muticopsis f. evoluta W.West & G.S.West (1911: 283, pi. 26, figs 121-123)

Navicula muticopsis f. reducta W.West & G.S.West (191 1 : 284, pi. 26, fig. 124)

Navicula muticopsiformis W.West & G.S.West (1911: 284, pi. 26, fig. 131)

Navicula muticopsis f. capitata Carlson (1913: 15, pi. 1, fig. 10)

Navicula mutica var. cymbelloides Perag. (Peragallo, 1921: 16, pi. 1, fig. 28)
Navicula mutica var. truncata Perag. (Peragallo, 1921: 17, pi. 1, fig. 27)
Navicula muticopsis var. capitata Perag. (Peragallo, 1921: 17, pi. 1, fig. 40)
See VanLandingham (1975: 2687) for details of synonymy.

additional references: Peragallo (1921: 17, pi. 1, fig. 27), Hustedt (1927-30: 614, fig. 1614),
Chalmers et al. ( 1 996: 40, pi. 12, figs e,

Fig. 2.86a-d

description: Cells solitary. Valves flat, slightly capitate in valve view; apical axis 10-25
um; transapical axis 6-10 um; pervalvar axis 4-6 um. Transapical striae radiating,
uniseriate, 10-14 in 10 um; areolae on the valve face rounded to slightly elongate; areolae on

153
 DIATOMS

the mantle solitary and rounded. Single stigma present near valve centre. Raphe-sternum
narrow; external central raphe endings and distal raphe endings strongly deflected in the
same direction away from the stigma. Girdle bands simple, each bearing a single row of
small poroids (Fig. 2.86d).

distribution: Coastal waters near Davis Station, East Antarctica; type locality, 65°15.05'S
64°30.0'E, Station 141, "Belgica" Expedition (van Heurck, 1909); Petermann I., Jenny I. and
Leonie I., Cape Tuxen (all Antarctic Peninsula) (Peragallo, 1921); South Shetland Is. and
South Orkney Is. (Hakasson & Jones, 1993).

Material collected from coastal waters off Davis Station showed all external raphe endings
(central and distal) deflected in the same direction, rather than in opposite directions as was
described for the genus (Mann, 1990).

Manguinea fusiformis (Manguin) Paddock (1988: 88, pi. 32)

BASIONYM: Tropidoneis fusiformis Manguin (1957: 130, pi. 6, fig. 39)
See Paddock (1990: 153) for additional synonyms.

ADDITIONAL REFERENCES: Manguin (I960: 322, pi. 16, figs 193a, b, 194, as T. fusiformis),
Paddock (1990: 153, pi. 19.1, fig. 14), Hasle & Syvertsen (1997: 289, pi. 65, tab. 72)

Fig. 2.67d

description: Cells usually lying in broad girdle view, fusiform, weakly silicified, lanceolate
in valve view; apical axis 49-135 um; pervalvar axis c. 11 um (at mid-point of cell 108 um
long). Raphe fin-like structures on narrow parts of valve. Transapical striae fine, aligned
somewhat diagonally, 26-28 in 10 um. Helictoglossae enlarged.
distribution: Antarctic Ocean (Paddock, 1990); Weddell-Scotia Confluence (Garrison et
ai, 1987, as Tropidoneis fusiformis); Terre Adelie (Manguin, 1957).

Membraneis challenged (Grunow) Paddock (1988: 81, pi. 29)

BASIONYM: Navicula challengeri Grunow, in Cleve & Grunow (1880: 64)
SYNONYM: Tropidoneis antarctica (Grunow) Cleve (1894: 24)

See Paddock (1990: 153) for additional synonyms.

additional REFERENCES: Hendey (1937: 350), Paddock (1990: 153, pi. 19.1, fig. 11), Hasle &
Syvertsen (1997: 287, pi. 64)

Fig. 2.83e

description: Cells usually united in ribbons; apical axis 85-270 um; pervalvar axis 33 um
(in cell with apical axis 100 um). Valve outline in girdle view smooth, lenticular. Raphe on
the raised valve surface; raphe fins located c. one-eighth of the apical axis from the valve
end. Transapical striae barely visible by LM, 18-24 in 10 um; central striae thickened,
varying in number and development. Helictoglossae enlarged.

distribution: Coastal waters off Davis Station, East Antarctica; Antarctic Ocean (Paddock,
1990); Weddell-Scotia Confluence (Garrison et al, 1987, as T. antarcticas [sic]).

Navicula criophila (Castrac.) De Toni (1891: 23)

BASIONYM: Pinnuiaria criophila Castrac. (Castracane, 1886: 26, pi. 15, fig. 2)

SYNONYMS: Navicula cryophila auct. non Manguin (1957: 127, pi. 5, fig. 30)
Navicula cryophila var. stauroptera auct. non Manguin (1957:127, pi. 5, fig. 31)
Navicula kryophila (see Poulin & Cardinal, 1982: fig. 38)

ADDITIONAL REFERENCES: Peragallo (1921: 56), Hustedt (1958: 145, figs 101-103)

description: [after Hustedt (1958)] Cells solitary, narrow, lanceolate; apical axis 28-60 um;
transapical axis6-10 um. Transapical striae parallel, uniformly spaced throughout valve, 6
in 10 um. Chloroplasts 2, extending the entire length of the cell.

54
 DIATOMS

Figure 2.86. Luticola muticopsis. a, SEM Valve face showing striae, raphe and ±central
solitary stigma (arrow), b, SEM One valve of a cell, c, SEM Other valve of the same cell as
b. d, SEM Cell in broad girdle view. Scale bars: a-d = 2 urn.

55
 DIATOMS

Figure 2.87. Navicula species, a-d, N. directa var. directa. a, SEM Solitary cell, b, SEM
Solitary cell, c, SEM Detail of central valve area, d, SEM Detail of terminal raphe ending, e.

f. V. cristata. e, SEM Solitary cell, f, SEM Detail of central raphe endings and transapical

striae. Scale bars: a, b, f = 2 urn; b, e = 10 urn.

156
 DIATOMS

distribution: Booth-Wandel I., Petermann I. (both Antarctic Peninsula) (Peragallo, 1921);

Weddell-Scotia Confluence (Garrison et al., 1987).

This uncommon species was not collected by us.

Navicula cristata Perag. (Peragallo, 1921: 56, pi. 2, fig. 1

1)

SYNONYMS: Navicula triplex VanLand. (VanLandingham, 1975: 2857))

Navicula subpolaris auct. non De Toni (1891), sensu Hendey (1937: 345)

Fig. 2.87e, f

description: Cells usually solitary, linear to lanceolate, with acute apices in valve view,
almost rectangular in girdle view; apical axis c. 100-105 urn; transapical axis c. 1-15 um. 1

Valve surface convex, with a prominent median raphe and a small hyaline central area;
central and terminal raphe endings small. Transapical striae c. 6-10 in 10 um.

distribution: Coastal waters off Davis Station, East Antarctica; Argentine Is., Antarctic
Peninsula (Peragallo, 1921); Bellingshausen Sea (Hendey, 1937).

Hendey (1937) described this as "a small characteristic Antarctic diatom observed in small
numbers in water obtained from melted ice".

Navicula directa (W.Sm.) Ralfs, in Pritchard (1861: 906) var. directa

BASIONYM: Pinnularia directa W.Sm. (Smith, 1853: 56, pi. 18, fig. 169)

See VanLandingham (1975: 2517) for further synonymy.

ADDITIONAL REFERENCES: Peragallo (1921: 56), Poulin & Cardinal (1982: 2837, fig. 10), Poulin
(1990: 138, pi. 18.1, fig. 24), Hasle & Syvertsen (1997: 280, pi. 63, tab. 70)

Fig. 2.87a-d

DESCRIPTION: Cells solitary, narrowly lanceolate, with acute to slightly rostrate apices; apical
axis 53-120 um; transapical axis 7-12 um. Transapical striae parallel, uniformly spaced, 7-
11 in 10 um, composed of slit-like poroids. Poroids 30-33 in 10 um, orientated
longitudinally in central part of valve, but obliquely towards the valve apices. Raphe central;
central raphe endings expanded into pores; terminal endings hooked in the same direction.
Chloroplasts 2, extending the entire length of cell, appressed to the girdle bands.

distribution: Coastal waters off Davis Station, East Antarctica; Southern Ocean, 57°35'S
105°46'E, offshore Wilkes Land; Arctic to Antarctic (Poulin, 1990); Ross Sea (Andreoli et
al., 1995); coastal waters off Syowa Station, East Antarctica (Ishikawa et al., 2001).

Navicula directa is reputedly a bottom-dwelling species that is found occasionally in the

plankton (Hendey, 1964). Navicula directa var. javanica Cleve, which also has an Arctic to
Antarctic distribution (Poulin, 1990), differs from the type variety in its larger size (apical
axis 86-145 um; transapical axis 11-12 um) and valve striation (cross-linear and parallel
striae 6-7 in 10 um).

Navicula gelida Grunow var. parvula Heiden, in Heiden & Kolbe (1928: 605, tab. 2,
fig. 53)

additional REFERENCE: Peragallo (1921: 56)

Fig. 2.88a-e

description: Cells solitary or joined together at valve surface, lanceolate; apical axis 21-27
um; transapical axis 5.6-6.5 um; pervalvar axis c. 2.9 um. Transapical striae 12-19 in 10
um, straight but slightly radiating towards the cell apices. Raphe central; endings expanded
into pores; terminal endings hooked in the same direction.

distribution: Coastal waters off Davis Station, East Antarctica; Antarctic coldwater species
(Kozlova, 1964).

157
 DIATOMS

Material collected near Davis Station consistently showed finer valve striation (striae 17-19
in 10 um) than previous descriptions (striae 12-15 in 10 urn; Heiden & Kolbe, 1928). The
less common N. gelida var. antarctica Heiden (Heiden & Kolbe, 1928) has larger cells
(apical axis 34-64 um; transapical axis 13-15 urn) with slightly coarser valve striation
(striae 8.5-11 in 10 urn).

Navicula glaciei van Heurck (1909: 1 1, pi. 1, fig. 13)

ADDITIONAL REFERENCES: Whitaker & Richardson (1980: 250), Poulin & Cardinal (1982: 2837,
fig. 28), Andreoli et al. (1995: 472, fig. 11)

Fig. 2.89a-e

DESCRIPTION: Cells solitary, linear to elliptical, with subrostrate apices; apical axis 29-42
um; transapical axis 7-13 um. Transapical striae 14-15 in 10 um, slightly radiating in
central area, straight nearer the apices. Raphe central; central raphe endings expanded (Fig.
2.89d); terminal endings hooked in the same direction (Fig. 2.89c). Axial area around raphe
narrow.

distribution: Coastal waters off Davis Station, East Antarctica; type locality, 65° 15.5'-
64°30.0'S, Antarctica (van Heurck, 1909); Scotia Sea, Antarctica (Poulin & Cardinal, 1982);
Ross Sea (Andreoli et al., 1995); coastal waters off Syowa Station, East Antarctica (Ishikawa
et al., 2001).

Navicula gracilis Ehrenb. var. antarctica Perag. (Peragallo, 1921: 57, pi. 2, fig. 14)

description: [after Peragallo (1921)] Cells solitary, lanceolate, with acute apices; apical axis
60-70 um. Transapical striae 10 in 10 um.
distribution: Petermann I., Antarctic Peninsula (Peragallo, 1921).

This rare variety was not seen by us.

Navicula schefterae Lobban (1986: 83, figs 2-10)

description: Cells forming monostromatic sacs with a short stalk, broadly

colonial,
elliptical; apical axis 25 um; transapical axis c. 10 um. Raphe straight, with apical endings
c.

bending weakly in the same direction. Striae weakly radiating throughout valve, 22-24 in 10
um, except in central area.
distribution: Type locality, Unwin Cove, Cape Legoupil, Antarctica (c. 63.3°S 58.0°W)
(Lobban, 1986).

Lobban (1986) noted that N. schefterae might be conspecific with N. cryophila and
(W.Sm.) Donkin, and that further SEM studies are required
N. crucicula to determine their
status.

Pinnularia quadratarea A. W.F.Schmidt var. constricta (0strup) Heiden (1905), in

Schmidt et al. (1874-1959: tab. 260, figs 3-8)
BASIONYM: Navicula pinnularia var. constricta 0strup (1895: 419, pi. 4, figs 23, 24)

ADDITIONAL REFERENCES: Heiden & Kolbe (1928: 596), Poulin (1990: 140, pi. 18.1, fig. 34)

Fig. 2.90a-c

description: Cells solitary. Valves lanceolate, but constricted at the centre; valve faces flat;
apical axis 35-96 um; transapical axis 11-19 um; pervalvar axis c. 4-14 um. Transapical
striae coarse, 10-1 in 10 um, biseriate (occasionaly triseriate). Raphe endings at valve poles
1

distinctly hooked, deflected in the same direction; central raphe endings expanded.
Chloroplasts 2, plate-like.

58
 * 1

DIATOMS

^m •
'
4
"•»!
;'''¥//''',',
"M
•ufJ
^Bf '• '
> • • • *

W/?; !!
¥ 1 , , M

'

1/: «,"
•-;-..>.

_d
Figure 2.88. Navicula gelida var. parvula. a, SEM Solitary cell in valve view, b, SEM
Oblique valve view, c, SEM Internal valve view, d, TEM Cells in girdle and valve view, e,
TEM Detail of valve striae and areolae. Scale bars: a-e = 2 urn.

159
 DIATOMS

Figure 2.89. Navicula glaciei. a, LM Cells in valve view, b, SEM Cell in valve view, c,
SEM Detail of terminal raphe ending, d, TEM Detail of central raphe endings, e, TEM Detail
of valve areolae. Scale bars: a = 10 (am; b = 5 (am; c-e =
1 (am.

160
 DIATOMS

Figure 2.90. Pinnularia quadratarea. a-c, P. quadratarea var. constricta. a, LM Cells in

valve and girdle view, b, SEM Cells in valve and girdle view, c, SEM Hooked raphe-end at
valve pole, d, P. quadratarea var. soederlundii, SEM Etched valve. Scale bars: a, b, d = 10
p.m; c = 1 urn.
 DIATOMS

Figure 2.91. Plagiotropus gaussii. a, LM Solitary cell in girdle view, b, SEM Solitary cell in
girdle view, c, TEM Solitary cell in girdle view, d, SEM Detail of cell end in girdle view, e,

TEM Detail of transapical striae and helictoglossae. Scale bars: a-c = 10 urn; d, e = 5 urn.

62
 DIATOMS

distribution: Sea-ice near Davis Station, East Antarctica; Arctic to Antarctic (Poulin,
1990); in pack-ice, "Gaussberg", Davis Sea (Heiden & Kolbe, 1928); coastal waters off
Syowa Station, East Antarctica (Ishikawa et al., 2001).
A single cell of P. quadratarea var. soederlundii Cleve was also collected in sea-ice near
Davis Station. This variety is smaller than var. constricta, being 35-66 urn long and with 1 1-
13 striae in 10 um. Var. soederlundii (Fig. 2.90d) was previously recorded off Heard I. and
the Davis Sea, East Antarctica by Heiden & Kolbe (1928: 595, tab. 2, figs 31, 32).

Plagiotropus gaussii (Heiden) Paddock (1988: 63, pi. 24)

BASIONYM: Tropidoneis gaussii Heiden, in Heiden & Kolbe (1928: 656, pi. 5, fig. 102)

ADDITIONAL REFERENCES: Paddock ( 1 990: 152, pi. 19.1, figs 5, 6), Hasle & Syvertsen (1997: 289,
pi. 65, tab. 72)

Fig. 2.91 a-e

description: Cells solitary or uniting into ribbons, often lying in girdle view. Valves
delicate, with a low gently curved ridge, resulting in the cell appearing bilobed in girdle
view; apical axis 102-159 um; pervalvar axis 20-35 urn. Transapical striae 15-18 in 10 urn.
Raphe fins absent (cf. Manguinea and Banquisea). Helictoglossae enlarged. Chloroplast 1,
multi-lobed, lying in the centre of the cell.

distribution: Coastal waters off Davis Station, East Antarctica; Antarctic, with ice
(Paddock, 1990); Lutzow-Holm Bay (Tanimura et al., 1990); Weddell-Scotia Confluence
(Garrison et al., 1987, as Tropidoneis gaussii); in pack-ice, "Gaussberg", Davis Sea (Heiden
& Kolbe, 1928, as Tropidoneis gaussii).
Several other species of Plagiotropus, Membraneis and Tropidoneis have been reported from
polar waters (for a fuller discussion and illustrations, see Paddock, 1988). Plagiotropus
australis (Perag.) Paddock is from Bransfield Strait (Paddock, 1988,
a rare species, reported
1990). Paddock stressed the need for further investigation of the "Tropidoneis" group of the
Naviculaceae.

Pleurosigma directum Grunow, in Cleve & Grunow (1880: 53)

See VanLandingham (1978: 3368) for details of synonymy.

ADDITIONAL REFERENCES: Hendey (1937: 348), Hasle & Syvertsen (1997: 282, pi. 62, tab. 71)

Fig. 2.92a-e

DESCRIPTION: Cells solitary. Valves flat, lanceolate, slightly sigmoidal, with subacute ends;
apical axis 180-270 um; transapical axis c. 44 um. Raphe straight or slightly sigmoidal;
central endings deflected in the same direction; pole endings deflected in opposite directions.
Valve striation fine, orientated in 3 distinct directions, one transverse and two oblique;
oblique striae 18.5 in 10 um.

distribution: Near Davis Station, East Antarctica; cosmopolitan (Hasle & Syvertsen, 1996);
near South Georgia (Hendey, 1937); coastal waters off Syowa Station, East Antarctica
(Ishikawa et al, 2001).

Hendey (1937) also reported P. smithianum Castrac. from Bransfield Strait. Published
descriptions of that rare species are rather incomplete.

Pseudogomphonema kamtschaticum (Grunow) Medlin, in Medlin & Round (1986:

216, figs 64-70)

BASIONYM: Gomphonema kamtschaticum Grunow (1878: 109(12), pi. 3, fig. 4)

ADDITIONAL REFERENCE: Medlin & Round (1990: 157, pi. 20.1, fig. 2)

Figs 2.93a-f, 2.94a, b

63
 DIATOMS

description: Cells solitary, clavate, sometimes with a short stalk, often lying in girdle view.
Valves linear to lanceolate, with rounded apices; apical axis 18-70 urn; transapical axis 4-11
urn; pervalvar axis at foot pole 2.9-3.1 urn, at head pole 8.5-10.5 urn. Transapical striae
slightly radiating throughout, 16-23 in 10 um (but somewhat fewer on primary valve),
uniseriate, containing slit-like poroids. Raphe central, present on both valves; central endings
expanded and pore-like; terminal endings hooked at foot pole, but reduced at head pole.
Cingulum composed of 4 non-porous bands, the 2 at the foot pole reduced to ligulae.
distribution: Near Davis Station, East Antarctica; Renwick Glacier; Prydz Bay.

Specimens collected near Davis on macroalgae or the underside of

Station, often epiphytic
the sea-ice, agree with the description provided by Medlin (Medlin & Round, 1986). Other
possible Antarctic varieties were also mentioned by Medlin (1990), and P. kamtschaticum
var. antarctica, intermediate between P. kamtschaticum and P. groenlandicum 0strup was
briefly described by Peragallo (1921). Medlin commented: "The varieties off. kamtschat-
icum will probably be described as separate species".

Trachyneis aspera (Ehrenb.) Cleve (1894: 191)

BASIONYM: Navicula aspera Ehrenb. (Ehrenberg, 1840: 213)
SYNONYMS: Stauroptera aspera (Ehrenb.) Ehrenb. (Ehrenberg, 1841: 135; 1843: 423, pi. 3, figs 3, 7)

Stauroneis aspera (Ehrenb.) Kiitz. (Kiitzing, 1844: 106, pi. 29, fig. 12)

Stauroneis achnanthes (Ehrenb.) Kiitz. (Kiitzing, 1844: 106, pi. 29, figs 20, 22)

Pinnularia aspera (Ehrenb.) Ehrenb. (Ehrenberg, 1854: pi. 19, fig. 26)
Navicula stauroneiformis Leud.-Fortm. (Leuduger-Fortmorel, 1892: 14)
Trachyneis aspera var. genuina Cleve (1894: 191), nom. inval.
Navicula aspera var. genuina (Cleve) Fricke (1902: 45), nom. inval.

See VanLandingham (1978: 4007) for further details of synonymy.

additional REFERENCES: Peragallo & Peragallo (1897-1908: 150, pi. 29, figs 1, 2), Hendey
(1937: 346, pi. 10, fig. 10), Proschkina-Lavrenko (1950: 230, pi. 79, fig. 1), Cleve-Euler (1955:
5), Hendey (1964: 236, pi. 29, fig. 13), Ricard (1987: 100, figs 834-839)

Fig. 2.95a-e

description: Cells solitary. Valves linear to elliptical, with rounded apices and deep
mantles; apical axis 180-220 um; transapical axis c. 40 um. Transapical striae radiating, 7-
10 in 10 um, uniseriate, composed of groups (alveoli) of c. 5 slit-like poroids; area
immediately around central raphe endings lacking poroids. Poroids 21-28 in 10 um, each
group opening internally by a single round hole (Fig. 2.95d). Raphe central; terminal raphe
endings hooked; central endings sinuous and deflected in the same direction as the terminal
endings (Fig. 2.95c). Chloroplasts 2, lying appressed to the girdle bands.

distribution: Near Casey Station, East Antarctica; Plough I., near Davis Station, East
Antarctica; Cape Tuxen, Petermann I., Port Lockroy, Argentine Is., King George I. (all
Antarctic Peninsula) (Peragallo, 1921); Scotia Arc at 53°44.5'S 39°29.5'W (Hendey, 1937);
Indian Ocean (Simonsen, 1974); Southern Ocean, 61°05'S 63°04'E and 69°06'S 78°21'E,
"Belgica" Expedition (van Heurck, 1909).

The slit-like poroids grouped into alveoli are a distinctive feature of this species. Some of the
specimens found in the current study were in the size range (apical axis 70-80 um) described
for T. aspera var. antarctica Perag. (Peragallo, 1921).

Family Auriculaceae

Auricula com pacta (Hust.) Medlin, in Medlin & Hasle (1990b: 473, fig. 14)

BASIONYM: Nitzschia compacta Hust. (Hustedt, 1958: 169, fig. 160)

ADDITIONAL REFERENCES: Paddock & Sims (1980: 161), Simonsen (1987: 450, pi. 670, fig. 1, as
N. compacta), Medlin & Hasle ( 1 990a: 1 59, pi. 2 1 .
1 ,
fig. 11)

64
 DIATOMS

Figure 2.92. Pleurosigma directum, a,LM Solitary cell in valve view, b, SEM Solitary cell
in valve view, c, SEM Solitary cell in oblique valve view, d, SEM Solitary cell, e, SEM
Detail of central raphe endings deflected in the same direction (arrows). Scale bars: a-d = 20
fim; e = 5 urn.

165
 DIATOMS

Figure 2.93. Pseudogomphonema kamtschaticiim. a, SEM Solitary cell in apical view, b,

SEM External head pole, c, SEM External foot pole, d, SEM Internal valve, e, SEM Internal
head pole, f, SEM Internal foot pole with pseudoseptum and apical chamber. Scale bars: a. d
= 5 um; b, c, e, f = 2 (im.

66
 DIATOMS

Figure 2.94. Pseudogomphonema and Auricula species, a, b, P. kamtschaticum. a, SEM Cell

in broad girdle view, b, SEM Cells in narrow girdle view, attached to macroalga. c-e,
A. compacta. c, SEM Cell in girdle (pervalvar) view, d, SEM Striae on valve, e, SEM Close
up of valve ends (arrows) in girdle view. Scale bars: a, b = 2 um; c = 10 urn; d, e = 1 uin.

67
 DIATOMS

Figure 2.95. Trachyneis aspera. a, SEMValve showing transapical striae, b, SEM Internal
valve view, c, SEM SEM Details of external endings of raphe, e,
Cell lying in valve view, d,
SEM Cell lying in girdle (pervalvar) view. Scale bars: a-c, e = 10 urn; d = 2 urn.

168
 DIATOMS

Figure 2.96. Cylindrotheca closterium. a, LM Straight cell, b, SEM Straight cell, c, SEM
Sigmoidal cell with rostrate ends, d, TEM Sickle-shaped cell with rostrate ends, e, SEM
Raphe subtended by cross bars (arrows) joining the narrow fibulae, f, TEM Raphe subtended
by cross bars (arrows) joining the narrow fibulae. Scale bars: a-d = 10 urn; e, f = um.
1

169
 DIATOMS

Fig. 2.94c-e

description: Cells solitary, asymmetrical, view) to auriculate or reniform

elliptical (girdle
(valve view); apical axis 79-134 um; transapical axis 12-13
um, with a curved strongly
ridged keel supporting the raphe and dividing the valve into a broad and narrow face. Fibulae
6-9 in 10 urn; striae 22-27 in 10 um, parallel in mid-region of valve, radiating outwards
towards the valve ends, containing double rows of small areolae. Central nodule present.
Copulae numerous, simple (Fig. 2.94e).

distribution: Offshore from Ranwick Glacier, Prydz Bay, East Antarctica; Inexpressible I.,

Ross Sea; type locality, 69°43'S 06°30'E (Hustedt, 1958); Antarctic sea-ice (Medlin &
Hasle, 1990b).

Nitzschia medioconstricta has a similar morphology to Auricula compacta, but the cells are
usually shorter and thinner and have a greater density of fibulae.

Family Bacillariaceae

Cylindrotheca closterium (Ehrenb.) Reimann & J.C.Lewin (1964: 289, pi. 124, figs 1-
4; pi. 125, figs 1-4)

BASIONYM: Ceratoneis closterium Ehrenb. (Ehrenberg, 1841: 144, pi. 4, fig. 7)

SYNONYMS: Nitzschia closterium (Ehrenb.) W.Sm. (Smith, 1853: 42, pi. 15, fig. 120), Hasle & Medlin (1990b:
177, pi. 23.1, figs 1-4)

Nitzschiella tenuirostris Mereschk. (Mereschkowsky, 1901: 476, pi. 4, figs 1-3)

See VanLandingham (1978: 3024) for further details of synonymy.

ADDITIONAL REFERENCES: Hasle (1964: 16, pi. 5, fig. 1; pi. 7, figs 1-12; pi. 8, figs 1-8; pi. 9, figs
1-9; pi. 10, figs 1-4), Hasle & Medlin (1990b: 177, pi. 23.1, figs 1-4), Moisan & Fryxell (1993:
493), Hasle & Syvertsen (1997: 294)

Figure 2.97. Diploneis stigmosa. a, SEM Cell in external valve view, b, SEM Slit-like
areolae on external valve surface, c, SEM Pore-like areolae on external valve surface. Scale
bars: a = 10 um; b, c = 1 um.

170
 DIATOMS

Fig. 2.96a-f

description: [after Hasle (1964) and Hasle & Syvertsen (1997)] Cells solitary, needle-like,
straight, sigmoidal or sickle-shaped, with valves sometimes twisted around the apical axis

and slightly rostrate ends. Valves weakly silicified; apical axis 30-400 urn; transapical axis
2.5-8 um. Fibulae narrow, 12-16 urn in 10 urn, fastened to the valve by short cross-bars
(Fig. 2.96e, f). Striae 70-100 in 10 um. Copulae numerous, narrow (Fig. 2.96e, f)- Central
nodule present.

distribution: Coastal waters off Davis Station, East Antarctica; near Drygalski Glacier,
Ross Sea; Southern Ocean, 57°26'S 1 12°08'E offshore Wilkes Land; cosmopolitan (Hasle &
Medlin, 1990); KERFIX, 50°40'S 68°25'E, Southern Ocean (Kopczyhska et al, 1998);
Liitzow-Holm Bay (Tanimura et al., 1990); Weddell-Scotia Confluence (Garrison et al.,
1987).

Using only LM observations, it is difficult to distinguish C. closterium from Nitzschia

longissima due to their similar size and shape. However, with EM, the raphe of
Cylindrotheca is seen to be interrupted in the centre, and the fibulae are narrow, arcuate and
fastened to the valve by short cross-bars. In N. longissima the fibulae are also narrow and
arcuate, but they join silicified ribs of the same diameter which run parallel to the raphe
fissure (see Hasle. 1964, pi. 11, fig. 2). Cylindrotheca is generally broader and is regarded as
the more delicate cell.

Diploneis stigmosa Heiden & Kolbe (1928: 613, pi. 2, fig. 56)

Figs 2.97a-c, 2.98a-d

description: Cells solitary, panduriform, with rounded poles in valve view; apical axis 58-
91 um; transapical axis 21-30 um. Striae 4.3-6 in 10 um, with radiating slits along each
margin and with a uniseriate row of pores along either side of raphe; pore size variable.
Raphe central; terminal fissures hooked externally (Fig. 2.98c). Internal longitudinal canals
present on either side of the raphe (Fig. 2.98d). Girdle consisting of a few open copulae (Fig.
2.98b).

distribution: Sea-ice and coastal waters near Davis Station, East Antarctica; Observatory
Bay, Kerguelen Is. (Heiden & Kolbe, 1928).

Specimens from Antarctic coastal waters near Davis Station measured 70-90 um long and
25-30 um wide (20-25 um at constriction), with striae 5-6 in 10 um.

Fragilariopsis curta (van Heurck) Hust. (Hustedt, 1958: 160, pi. 11, figs 140-144; pi.

12, fig. 159)

BASIONYM: Fragilaha curta van Heurck (1909: 24, pi. 3, fig. 37)

SYNONYMS: Fragilariopsis linearis var. curta (van Heurck) Freng., in Frenguelli & Orlando (1958: 107, pi. 4,

fig. 31)
Nitzschia curta (van Heurck) Hasle (1972a: 115), Buck & Garrison (1983: 1264, fig. 8), Everitt & Thomas
(1986: 9, fig. 2E), Fryxell (1989: 12)

See Hasle (1965b) for full synonymy, LM and EM descriptions, discussion of nomenclatural problems and
distribution.

ADDITIONAL REFERENCES: Hendey (1937: 330, as Fragilaha curta), Hasle (1965b: 32, pi. 6, fig.
6; pi. 12, figs 2-5; pi. 13, figs 1-6; pi. 16, fig. 6; pi. 17, fig. 5), Hasle & Medlin (1990c: 181, pi.
24.6, figs 2-5), Moisan & Fryxell (1993: 493)

Fig. 2.99a-f

description: Cells solitary or chain-forming, attached by the valve surfaces, ±linear in valve
view, with rounded poles, one broader than the other; apical axis 10-42 um; transapical axis

171
 DIATOMS

3.5-6 urn. Transapical striae 9-12 in 10 um, straight, but striae near valve poles curved.
Interstrial membranes perforated by (1-) 2 (-3) rows of areolae, c. 30 in 10 um. Fibulae
present in slightly lower numbers than striae, comparatively broad and irregularly placed.
Pseudonodulus absent. Terminal nodules not confirmed (Hasle, 1965b).

distribution: Drygalski Glacier, Ross Sea; type locality, dark greenish brown sea-ice,
65°15.5'S, 64°30.0'W; KERFIX, 50°40'S 68°25'E, Southern Ocean (Kopczyhska et al,
1998); sea-ice near Davis Station (Archer et al., 1996b); BRATEGG samples from the
Antarctic (Hasle, 1965b); 74°40'-77°37'S (Cassie, 1963); Antarctic ice-edge to the mid-
most commonly observed in the Antarctic (Hustedt,
tropical zone of the South Atlantic, but
1958); Weddell-Scotia Confluence (Garrison et al., 1987); coastal waters off Syowa Station,
East Antarctica (Ishikawa et al, 2001).

Hendey (1937) described being characteristic of the coastal diatom flora of

this species as
land masses within the Polar Front. Manguin (1960) observed F. curta at all sampling
stations within the Antarctic zone and in greatest numbers near the coast, as did Kozlova
(1962). Due to its size range and general shape, F. curta is easily confused with F. cylindrus;
the former is slightly heteropolar and with striae 9-12 in 10 jj.m, whereas F. cylindrus is
isopolar (linear) with striae 13-17 in 10 um.

Fragilariopsis cylindrus (Grunow) Willi Krieg., in Helmke & Krieger (1954: 17, pi.
187-189)
BASIONYM: Fragilaha cylindrus Grunow, in Cleve (1883: 484, pi. 37, fig. 64a-c)

SYNONYMS: Fragilaria nana Steem. Niels. (Steemann Nielsen, 1935: 74, fig. 2a, b)

Fragilariopsis cylindrus var. planctonica Willi Krieg., in Helmske & Krieger (1954: 17, pi. 188, 189)

Fragilariopsis cylindrus f. minor Manguin (1960: 298, pi. 12, fig. 127)
Fragilariopsis linearis (Castrac.) Hust. var. intermedia Manguin (1960: 298, pi. 2, fig. 129)

Fragilariopsis nana (Steem. Niels.) Paasche (1961: 201) pro parte

Nitzschia cylindrus (Grunow) Hasle (1964: pi. 3, fig. 3)

See Hasle (1965b) for full synonymy, LM and EM, descriptions, discussion of nomenclatural problems and
distribution.

ADDITIONAL REFERENCES: Buck & Garrison (1983: 1264, fig. 7), Hasle & Medlin (1990c: 181, pi.
24.6, figs 6-12), Moisan & Fryxell (1993: 493), Hasle & Syvertsen (1997: 302, pi. 68)

Fig. 2.100a-e

description: Cells solitary or chain-forming, attached by the valve surfaces, linear in valve
view, with rounded ends; apical axis 3-48 um; transapical axis 2-4 um. Transapical striae
13-17 in 10 jam, straight. Interstrial membranes perforated by minute areolae, either
scattered or in 2 or 3 (exceptionally 4) rows, 60-70 in 10 um. Fibulae present as
continuations of the costae. Canal raphe present.

distribution: Sea-ice near Davis Station, East Antarctica (Archer et al., 1996b); type
locality, Tindingen, Greenland, Cape Wiinkerema near Bering Strait (Cleve, 1883); KERFIX,
50°40'S 68°25'E, Southern Ocean (Kopczyhska et al., 1998); Zemlya Frantsa-Iosifa (Franz
Josef Land), Arctic Ocean (Grunow, 1884); bipolar, cold-water species mainly confined to
Antarctic and Arctic waters (Hasle, 1965b); Weddell Sea (Buck & Garrison, 1983); Weddell-
Scotia Confluence (Garrison et al., 1987); Benguela Current (Heiden & Kolbe, 1928).

Hasle (1965b) explained that when Paasche transferred Fragilaria nana Steem. Neils, to
nana with some reservation, suspecting that more than
Fragilariopsis, he retained the epithet
one species had been identified as Fragilaria nana in the Norwegian phytoplankton studies
of the 1950s and 1960s. The diagnosis of Fragilariopsis nana (Steem. Neils.) Paasche
apparently included the linear-valved Fragilaria nana Steem. Neils. Fragilariopsis nana is
now separated into two taxa, viz. the linear-valved Fragilaria nana (a synonym of
Fragilariopsis cylindrus) and the elliptical-lanceolate-valved Fragilariopsis pseudonana
Hasle.

172
 DIATOMS

Figure 2.98. Diploneis stigmosa. a, SEM Cell in external valve view, b, SEM Cell in broad
girdle view, c, SEM Close up of valve with hooked raphe end. d, SEM Internal longitudinal

canals. Scale bars: a-d = 10 urn.

73
 DIATOMS

Figure 2.99. Fragilariopsis carta, a. SEM Solitary cell, valve view. b. SEM Internal valve
view, c, SEM Detail of areolate striae, d, SEM Two cells in girdle view, e, SEM Two cells in

oblique view, f, TEM Valve view showing distinct striae and areolae. Scale bars: a-f = 2 urn.

174
 DIATOMS

Figure 2.100. Fragilariopsis cylindrus. a, SEM Two cells in girdle view, b, TEM Cell with
details of fibulae, striae
and areolae, c, SEM Cell in apical view, d, TEM Detail of areolate
striae, e, SEM Internal valve view. Scale bars: a-e = 1 urn.

175
 DIATOMS

Figure 2.101. Fragilariopsis kerguelensis. a, LM Solitary cell, b, SEM Etched valve of a

solitary cell. c. SEM Etched valve of a solitary cell, d, SEM Oblique valve view, e, SEM
Internal valve view, f, SEM Chain of cells lying in girdle (pervalvar) view. Scale bars: a. b,
e, f = 5 urn; c, d =1 um.

176
 DIATOMS

/p^

\&

Figure 2.102. Fragilariopsis species, a, b, F. linearis, a, LM Solitary cells in valve and

girdle view, b, LM Cell in valve view, c-e, F. obliquecostata. c, LM Cell in valve view, d,
SEM Cell in valve view, e, SEM Detail of striae. Scale bars: a-e = 10 urn.

77
 DIATOMS

Figure 2.103. Fragilariopsis pseudonana. a, SEM Solitary cell in oblique valve view, b,
SEM Valve view with extreme eccentric canal raphe (arrows), c. SEM Chain of cells lying in
narrow girdle view. d. SEM Cell in broad girdle view. Scale bars: a-d = 2 um.

7S
 )

DIATOMS

Fragilariopsis kerguelensis (O'Meara) Hust. (Hustedt, 1952: 294)

BASIONYM: Terebraria kerguelensis O'Meara (1877: 56, pi. 1, fig. 4)

SYNONYMS: Fragilaria antarctiea Castrac. (Castracane, 1886: 56. pi. 25. Tig. 12)

Trachysphenia australis var. kerguelensis (O'Meara) De Toni (1892: 645)

Fragilariopsis antarctiea (Castrac.) Hust., in Schmidt (1913: pi. 299, figs 9-14)

Nitzschia kerguelensis (O'Meara) Hasle (1965b: 14, pi. 3, figs 4, 5; pi. 4, figs 1 1-18; pi. 5, figs 1-1 1; pi. 6,

figs 2-4; pi. 7, fig. 9; pi. 8, fig. 10; pi. 16, figs 3-5), Hasle (1972a: 115), Hasle & Medlin (1990c: 181. pi.

24.2, figs 11-18; pi. 24.3, fig. 9), Moisan & Fryxell (1993: 493)

See Hasle (1965b) for full synonymy, LM and EM descriptions, discussion of nomenclatural problems and
distribution.

ADDITIONAL REFERENCES: Hustedt (1958: 162, figs 121-127), Hasle & Syvertsen (1997: 296, pi.

66)

Fig. 2.101a-f

description: solitary or forming ribbon-shaped colonies attached by the valve

Cells
surfaces, in view valve face slightly curved towards ends, in valve view cells
girdle
elliptical; apical axis 10-76 urn; transapical axis 5-11 urn. Transapical striae comparatively
coarse, 4-7 in 10 urn, straight, those near the valve poles slightly curved. Interstrial
membranes punctate, with 2 rows of alternating areolae, 8-10 in 10 um.
distribution: Southern Ocean, 57°35'S 105°46'E, offshore Wilkes Land; sea-ice near Davis
Station (Archer el ai, 1996b); type locality, surface of the water between Kerguelen and
Heard Islands (O'Meara, 1877); KERFIX, 50°40'S 68°25'E, Southern Ocean (Kopczyhska et
ai, 1998); circumpolar Antarctic distribution (Baker, 1954); Weddell-Scotia Confluence
(Garrison et ai, 1987).

This species has been described as "the most abundant diatom in Antarctic Seas" (Hart,
1942).

Fragilariopsis linearis (Castrac.) Freng. (Frenguelli, 1943: 244, pi. 1, fig. 16)

BASIONYM: Fragilaria linearis Castrac. (Castracane, 1886: 56, pi. 19, fig. 9; pi. 25, fig. 1 1

SYNONYM: Nitzschia lineata (Castrac.) Hasle (1972a: 1 15)

See Hasle (1965b) for full synonymy and description.

ADDITIONAL REFERENCES: Heiden & Kolbe (1928: 550, pi. 6, fig. 128), Hustedt (1958: 161, pi.
11, figs 137-139; pi. 12, fig. 152), Hasle (1965b: 37, pi. 12, fig. 17; pi. 15, figs 9-11), Hasle &
Medlin (1990c: 181, pi. 24.6, fig. 17; pi. 24.7, figs 9-11)

Fig. 2.102a, b

description: Cells solitary, linear in valve view, with obtusely rounded poles; apical axis
40-72 um; transapical axis 7-9 um. Transapical striae 7.5-9 in 10 um, usually straight, but
sometimes curved at the poles. Interstrial membranes punctate, with 2 rows of areolae c. 25
in 10 um. Canal raphe markedly eccentric. Fibulae 7.5-9 in 10 um. Central nodule absent.

distribution: Coastal waters off Davis Station, East Antarctica; BRATEGG Stations 1 1, 43,
44, 47, 49, 54, 56 (Bellingshausen Sea-Amundsen Sea) (Hasle, 1965b); Weddell-Scotia
Confluence (Garrison et ai, 1987; Moisan & Fryxell, 1993).

Fragilariopsis obliquecostata (van Heurck) Heiden, in Heiden & Kolbe (1928: 555)
BASIONYM: Fragilaria obliquecostata van Heurck (1909: 25, pi. 3, fig. 38)
SYNONYMS: Fragilaria antarctiea van Heurck (1909: 25, pi. 3, fig. 38)
Fragilaria obliquecostata f. maxima van Heurck (1909: 25, pi. 3, fig. 40)
Fragilaria antarctiea f. lata van Heurck (1909: 25, pi. 3, fig. 4)

79
 DIATOMS

Fragilaria sublinearis van Heurck (1909: 25, pi. 3, fig. 37)

Fragilariopsis sublinearis (van Heurck) Heiden & Kolbe, in Frenguelli (1943: 241, pi. 1, figs 12?, 13-15)
Nitzschia obliquecostata (van Heurck) Hasle (see Hasle & Medlin, 1990c: 182, pi. 24.3, figs 2-7)

See Hasle (1965b) for full synonymy, LM and EM descriptions and discussion of nomenclatural problems.

ADDITIONAL REFERENCES: Hustedt (1958: 163, pi. 12, figs 149-151), Manguin (1960: 299, pi. 13,
figs 136, 137; pi. 29, fig. 354), Fukushima (1962, pi. 2, figs 12, 13), Jouse et al. (1962: 86, pi. 3,
fig. 9), Hasle (1965b: 18, pi. 7, figs 2-7), Hargraves (1968: 60, figs 103, 104)

Fig. 2.102c-e

description: Cells solitary, linear to lanceolate in valve view; apical axis 57-110 urn;
transapical axis8-10 urn. Transapical striae 6.5-8 in 10 um, usually straight, but sometimes
curved or oblique near the poles. Interstrial membranes punctate, with 2 or 3 rows of areolae;
areolae 16-23 in 10 um. Canal raphe markedly eccentric. Fibulae 6.5-8 in 10 um. Central
nodule absent.

distribution: Southern Ocean and Antarctic coastal waters near Davis Station, East
brown sea-ice, 65°15.5'S 64°30.0'W (van Heurck,
Antarctica; type locality, dark greenish
1909); Weddell-Scotia Confluence (Garrison et al, 1987).

Specimens from coastal East Antarctica seen by us have slightly less dense areolae (16-20 in
10 jam). Heiden & Kolbe (1928) reported 20, and Manguin (1960) 23 areolae (poroids) in 10
um (Hasle, 1965b).

Fragilariopsis pseudonana (Hasle) Hasle (1993: 317)

BASIONYM: Nitzschia pseudonana Hasle (1974: 427), Hasle & Medlin (1990c: 181, pi. 24.1, figs 7-14;
pi. 24.2, figs 20, 21; pi. 24.8, figs 1-3)

SYNONYM: Fragilaria nana Steem. Neils., in Holmes (1956: fig. Ma, pro parte), Paasche (1961: 201, pi. la,
fig. la), Hasle (1964: pi. 3, fig. 2)

See Hasle (1965b) for full synonymy, LM and EM descriptions and discussion of nomenclatural problems.

additional REFERENCES: Hasle (1965b: 22: pi. 1, figs 7-14; pi. 4, figs 20, 21; pi. 8, figs 1-9;
pi. 17, fig. 6), Moisan & Fryxell (1993: 493), Kang & Lee (1995: 260, fig. 6C), Hasle & Syvertsen
(1997: 300, pi. 67)

Fig. 2.103a-d

description: Cells solitary or in short chains with cells attached by valve faces, elliptical to
lanceolate in valve view; larger valves with slightly elongated ends; apical axis 4-20 um;
transapical axis 2.5-5 um. Transapical striae straight in the middle of the cell, curved near
the valve poles, 18-28 in 10 um. Canal raphe markedly eccentric (Fig. 2.103b). Fibulae
present in the same number as striae, arcuate, fastened to valve near striae and valve mantle;
no indication of a greater distance between the 2 fibulae in the middle. Interstrial membranes
perforated by 2 rows of minute areolae, 6-8 in 10 um, situated close to striae. Outer canal
wall without areolae. Pseudonodulus absent; terminal nodules present.

distribution: Southern Ocean, south of Australia; Drygalski Glacier, Ross Sea; BRATEGG
sample, brownish undersurface of pack-ice, 62°30'S, 19°42'W (Hasle, 1965); KERFIX,
50°40'S 68°25'E, Southern Ocean (Kopczyhska et al, 1998); Bransfield Strait (Kang & Lee,
1995); Weddell-Scotia Confluence (Garrison et al, 1987).

Fragilariopsis pseudonana is also known from cold northern waters (Denmark Strait, 62°N
33°W and 60°18'N 31°00'W; 61°08.5'N 02°21'W) as well as warmer regions
Shetland Is.,

(Gulf of California, Gulf of Panama, Wellington Harbour; Hasle, 1965b). Due to the
similarity of cell shape and striation, it is difficult to separate F. pseudonana from small
specimens of F. cylindrus. See discussion under the latter species.

180
 DIATOMS

Figure 2.104. Fragilariopsis rhombica. a, LM Cell in valve view, external, b, SEM Cell in
valve view, c, SEM External valve view, d, SEM Internal valve view. Scale bars: a-d = 2 \im.
 DIATOMS

Figure 2.105. Fragilariopsis ritscheri. a, LM Solitary cell in valve view, b, SEM Solitary
cell in valve view, c, SEM Solitary cell in valve view, d, SEM Interstrial poroids aligned in
alternate rows. Scale bars: a-c = 10 \im; d = 1 urn.

182
 DIATOMS

Fragilariopsis rhombica (O'Meara) Hust. (Hustedt, 1952: 296, figs 6, 7)

BASIONYM: Diatoma rhombicum O'Meara (1877: 22, pi. 1, fig. 2)

SYNONYMS: Fragilaria rhombica (O'Meara) Heiden & Kolbe (1928: 552, pi. 6, figs 129, 130)

Nitzschia antarctica Okuno ( 1954: 24, pi. 3, fig. 7)

Nitzschia angulata Hasle (1972a: 115)

ADDITIONAL REFERENCES: Hustedt (1958: 163, figs 113-120), Hasle (1965b: 24, pi. 1, fig. 6; pi.

4, fig. 19; pi. 6, fig. 5: pi. 8, fig. 11; pi. 9, figs 1-6; pi. 10, figs 2-6), Hasle & Medlin (1990c: 181,
pi. 24.1, fig. 6; pi. 24.4, figs 1-6), Moisan & Fryxell (1993: 493), Hasle & Syvertsen (1997: 300,
pi. 68, tab. 73)

Fig. 2.104a-d

description: Cells circular to elliptical, pointed towards the poles; apical axis 8-53 urn;
transapical axis 7-13 urn. Transapical striae straight in the centre, curved towards the poles,
8-16 in 10 um, somewhat thickened internally (Fig. 2.104c). Interstrial membranes
comprising (1 or) 2 rows of poroid areolae; areolae 22-26 in 10 um. Fibulae 8-16 in 10 um.
Central nodule absent.

distribution: Coastal waters near Davis Station, past Antarctica; Terra Nova Bay; Southern
Ocean, 57°35'S 105°46'E, offshore Wilkes Land; Weddell-Scotia Confluence (Garrison et
al, 1987); BRATEGG Antarctic samples (Hasle, 1965b); Heard I. (Manguin, 1954); Indian
Ocean sector of the Antarctic (Jouse et al., 1962; Kozlova, 1962); Bellingshausen Sea and
South Orkneys (Frenguelli & Orlando, 1958).

Cells found near Davis Station have an apical axis of 23-36 um, striae (10-) 12-15 in 10
um, fibulae 12-15 in 10 um and areolae 26 in 10 urn.

Fragilariopsis rhombica, F. separanda and Nitzschia barbieri are rather similar (Table 2.1).
Hustedt (1958) regarded the valve outline of F. rhombica to be predominantly lanceolate to
linear, while that of F. separanda was linear. However, illustrations of the two species in
Hasle (1965b) show considerable overlap of valve shapes.

Table 2.1. Comparison of Fragilariopsis rhombica, F. separanda and Nitzschia barbieri

(data from Hasle & Medlin, 1990c).

Apical axis (um) Striae in 10 um Rows of areolae

F. rhombica 8-53 8-16 1-2

F. separanda 10-33 10-14 1

N. barbieri 49-118 8-11 1 (-2)

Fragilariopsis ritscheri Hust. (Hustedt, 1958: 164, pi. 11, figs 133-136; pi. 12, fig.
153)
SYNONYMS: Nitzschia ritscheri (Hust.) Hasle (1972a: 115), Hasle & Medlin (1990c: 181, pi. 24.1, fig. 20; pi.
24.2, figs 1-10; pi. 24.3, fig. 8)

IFragilaria sublinearis var. ambigua Perag. (Peragallo, 1921 : 68, pi. 3, fig. 4)

ADDITIONAL REFERENCES: Hasle (1965b: 20, pi. 1, fig. 20; pi. 3, fig. 3; pi. 4, figs 1-10; pi. 5, figs

12, 13; pi. 6, fig. 1; pi. 7, fig. 8), Jouse et al. (1962: 86, pi. 6, fig. 9), Kozlova (1962: pi. 9, fig.

11)

Fig. 2.105a-d

DESCRIPTION: Cells solitary or in chains; small cells linear to elliptical in valve view; large
cells linear to lanceolate and broader at one pole; apical axis 22-60 um; transapical axis 8-9

183
 DIATOMS

um. Transapical striae straight in the middle of the cell, curved near the valve poles; striae
adjacent to broad more oblique than at narrow pole, 6-8 in 10 urn. Interstrial
pole
membranes perforated by 2 or 3 rows of usually alternating poroids, 18-24 in 10 um. Fibulae
curved, present in the same number as striae. Pseudonodulus absent. Canal raphe present, but
not always obvious.

distribution: Southern Ocean and coastal Antarctica, south of Australia; Southern Ocean,
57°35'S 105°46'E, offshore Wilkes Land, also 60°50'S 93°36'E, offshore Wilhelm II Land;
type locality 69°43'S, 06°30'E to 61°32'S, 0°00' (Hustedt, 1958); sub-Antarctic and
Antarctic (Hustedt, 1958; Kozlova, 1962; Hasle, 1965b); Weddell-Scotia Confluence
(Garrison et al., 1987); Liitzow-Holm Bay (Tanimura et al, 1990).

Specimens collected by us in the Southern Ocean and coastal Antarctica, south of Australia,
were of solitary cells, some being a little longer (60 um) than previously recorded.

Fragilariopsis separanda Hust. (Hustedt, 1958: 165, pi. 10, figs 108-1 12)

SYNONYM: Nitzschia separanda (Hust.) Hasle (1972a: 115), Hasle & Medlin (1990c: 181, pi. 24.4, figs 7-10)

See Hasle (1965b) for full synonymy, LM and EM descriptions and discussion of nomenclatural problems.

ADDITIONAL REFERENCES: Jouse et al. (1962: 86, pi. 3, fig. 8), Kozlova (1962: pi. 9, fig. 10),
Hasle (1965b: 26, pi. 9, figs 7-10; pi. 10, fig. 1), Hasle & Syvertsen (1997: 302, pi. 68, tab. 73)

description: [after Hasle (1965b)] Cells solitary, elliptical to lanceolate in valve view when
small, broad and linear with tapering ends when large; apical axis 10-33 um; transapical axis
8-13 um. Transapical striae 10-14 in 10 um. Interstrial membranes perforated by 1 row (2
rows near the margin) of short tube-like poroids, 12-16 in 10 um. Canal raphe with broad
fibulae and narrow apertures between them. Raphe ends with a minute notch.

distribution: South Atlantic-Antarctic, 51°29'S 0°08'W to 69°43'S 06°30'E (Hustedt,

1958); Antarctic, BRATEGG (Hasle, 1965b); Liitzow-Holm Bay (Tanimura et al., 1990).

While we have not seen this species, F, separanda has been compared with F. rhombica
(angulata), probably the most closely related taxon (Hustedt, 1958). In at least some
investigations of Southern Ocean planktonic diatoms, F. separanda is likely to have been
misidentified as F. rhombica (Hasle, 1965b).

Fragilariopsis sublinearis (van Heurck) Heiden, in Heiden & Kolbe (1928: 554)
BASIONYM: Fragilaria sublinearis van Heurck (1909: 35, pi. 3, fig. 39)
SYNONYMS: Fragilaria sublinearis f. longa van Heurck (1909: 35, pi. 3, fig. 41)
Nitzschia sublineata Hasle (1972a: 115), Hasle & Medlin (1990c: 181, pi. 24.3, fig. 1; pi. 24.5, figs 1-10; pi.

24.6, fig. 1)

Fragilaria sublinearis var. A., fide Bunt (1961: 32, pi. 8)

See Hasle (1965b) for full synonymy, LM and EM descriptions, distribution and discussion of nomenclatural
problems.

additional references: Jouse et al. (1962: 87, pi. 6, figs 10-16), Hasle (1965b: 27, pi. 7, fig.

1; pi. 11, figs 1-10; pi. 12, fig. 1; pi. 17, fig. 1), Moisan & Fryxell (1993: 493)

description: Cells solitary, narrowly linear to lanceolate, with obtuse apices in valve view;
apical axis 30-95 um; transapical axis 5.5-8 um. Transapical striae 7.5-9 in 10 um, broad
and straight, 1-3 striae near valve poles oblique or curved. Interstrial membrane perforations
barely visible, comprising 2 rows of minute alternating poroids, 35-39 in 10 um. Fibulae
present in the same number as striae. Keel with puncta of irregular width. Pseudonodulus
absent.

distribution: Type locality, dark greenish brown sea-ice, 65°15.5'S, 64°30.0'W; Antarctica
(Heiden & Kolbe, 1928); brownish undersurface of shelf-ice, 78°00'S 38°45'W, net samples
from BRATEGG Station 49, near Peter I Island and Station 57 at the pack-ice (Hasle,
1965b); Liitzow-Holm Bay (Tanimura et al., 1990); South Georgia (Hendey, 1937); coastal
waters off Mawson Station, East Antarctica (Bunt, 1961); Indian Ocean sector of the
Antarctic (Kopczyhska et al., 1986); Terre Adelie (Frenguelli, 1960).

184
 DIATOMS

Fragilariopsis vanheurckii (Perag.) Hust. (Hustedt, 1958: 166, pi. 12, figs 154-156)
BASIONYM: Fragilaria vanheurckii Perag. (Peragallo, 1921: 68, pi. 3, fig. 9, as van Heurckii)

SYNONYMS: Nitzschia vanheurckii (Perag.) Hasle (1972a: 1 15)

IFragilaria cylindrus f. elongala van Heurck (1909: pi. 3, fig. 43)

See Hasle (1965b) for full synonymy, LM and EM descriptions, distribution and discussion of nomenclatural
problems.

ADDITIONAL REFERENCES: Hasle (1965b: 30, pi. 12, figs 13-16; pi. 13, figs 7, 8; pi. 15, fig. 8),
Hasle & Medlin (1990c: 181, pi. 24.6, figs 13-16; pi. 24.7, fig. 8)

Fig. 2.106a-c

description: Cells solitary or in ribbon-shaped colonies, linear in valve view, with

extremely short broadly rounded poles; apical axis 35-77 urn; transapical axis c. 5 urn.
Transapical striae strongly silicified, 10-11 in 10 urn, straight; striae near poles slightly
curved or oblique; striae visible in girdle view. A few short apical ribs arising from striae
nearest the poles. Interstrial membrane perforations barely visible, comprising 2 rows of
minute close-set poroids, 42-51 in 10 urn. Fibulae 7-9 in 10 urn; apertures between fibulae
mostly narrow; central aperture larger. Stria adjacent to large central aperture shorter than
others. Outer fissure of canal raphe visible in parts of the valve, especially near one pole.
Terminal nodules and an interruption of the fissure opposite the large middle aperture.
Poroids lacking in the outer canal wall.

DISTRIBUTION: Southern Ocean, south of Australia; type locality, Argentine Is., Antarctic
Peninsula (Peragallo, 1921); BRATEGG sample at 69°12'S 94°06'W, also brownish
undersurface of shelf-ice, 78°00'S 38°45'W (Hasle, 1965b); Weddell-Scotia Confluence
(Garrison et ah, 1987); Liitzow-Holm Bay (Tanimura et al., 1990).

Nitzschia acicularis (Kiitz.) W.Sm. (Smith, 1856: 43, pi. 15, fig. 122)

BASIONYM: Synedra acicularis Kiitz. (Kiitzing, 1844: 63, pi. 4, fig. 3)

See VanLandingham (1978: 3002) for details of synonymy.

Figure 2.106, Fragilariopsis vanheurckii. a, SEM Cell in valve view, b, SEM Detail of
transapical striae, c, SEM Cell in valve view. Scale bars: a, c = 10 p.m; b = 1 |im.

185
 DIATOMS

Figure 2.107. Nitzschia species, a-c, N. acicularis. a, TEM Single cell in valve view,
shadow-cast preparation, b, SEM Cell in oblique valve/girdle view. c. SEM Cell in broad
girdle view, d-f, N. australis. d, SEM Cell in oblique valve/girdle view, e, SEM Cell in
broad girdle view, f, SEM Cell in girdle view with numerous copulae. Scale bars: a, b = 2
um; c = 1 urn; d-f = 10 urn.

186
 DIATOMS

ADDITIONAL REFERENCES: Krammer & Lange-Bertalot (1988: 123), Chalmers et al. (1996: 45, pi.

15a)

Fig. 2.107a-c

DESCRIPTION: Cells solitary, lanceolate in apical view, with rostrate ends; apical axis 22-150
|im; transapical axis 1.5-5 um. Transapical striae straight, close-set, 40-50 in 10 urn. Fibulae
15-27 in 10 \\.m. Central nodule absent.

distribution: Southern Ocean, 63°26'S 99°39'E, offshore Shackelton Ice Shelf; Signy I.,

South Orkneys (Chalmers et al., 1996).

Nitzschia acicularis is morphologically similar to N. decipiens (see below). The specimens

collected by us included cells of dimensions 22-25 um apical axis, 1.5-2.5 um transapical
axis, striae 40-50 and fibulae 20-27 in 10 um.

Nitzschia australis (Perag.) A.Mann (1937: 63, pi. 6, fig. 1)

BASIONYM: Nitzschia dubia var. australis Perag. (Peragallo, 1921: 65, pi. 3, figs 17, 18)

SYNONYM: Pseudonitzschia australis auct. non Frenguelli (1939), sensu Hustedt (1955: 43, pi. 15, fig. 1).

See Hasle (1994: 1037)

Fig. 2.107d-f

description: Cells solitary, biarcuate and rostrate; apical axis 83-110 um; transapical axis
10-15 um. Valves with a curved strongly ridged keel dividing them into 2 faces. Transapical
striae straight, 19-25 in 10 um, each composed of 2 rows of areolae. Central nodule present.
Copulae numerous, simple (Fig. 2. 1 07f).

distribution: Coastal waters off Davis Station, East Antarctica; Petermann I., Antarctic
Peninsula (Peragallo, 1921); Antarctic (Mann, 1937).

According to VanLandingham (1967: 3038), N. dubia var. australis is the valid name for
N. australis. However, we agree with Mann (1937) in retaining N. australis as a separate
species. Nitzschia hybrida Grunow, N. bilobata W.Sm., N. dubia W.Sm. and N. amphiprora
(Cleve) Grunow are all rather similar, but they do not resemble N. australis.

Specimens reported by Mann (1937) were 98-1 10 um long, with the transapical axis 12-15 um
and 20 striae in 10 urn. Those of Peragallo (1921) were 90-100 um long, with the transapical
axis 10-12 um and 19-20 striae in 10 um. Cells collected by us measured 83-98 u m in
length, with 25 striae in 10 um.

Nitzschia barbieri Perag. (1921: 66)

SYNONYM: Nitzschia ostenfeldii van Heurck var. minor van Heurck (1909: 22, pi. 3, fig. 177)

ADDITIONAL REFERENCES: Hasle (1965a: 41, pi. 15, figs 6, 7), Hasle & Medlin (1990c: 182,
pi. 24.7, figs 6, 7; pi. 24.4)

description: Cells solitary, in apical view elongate with parallel sides pointed towards the
poles; apical axis 49-118 um; transapical axis 8-14 um. Transapical striae straight in the
centre, curved towards the poles, 8-11 in 10 um. Interstrial membranes comprising
1 (occasionally 2) rows of poroid areolae; areolae c. 15 in 10 u Fibulae 8-11 in 10 um.m -

Central nodule absent.

distribution: Type locality, Argentine Is. and Petermann I., Antarctic Peninsula (Peragallo,
1921); southern BRATEGG stations, Bellingshausen Sea (Hasle, 1965a); Antarctic (Hasle &
Medlin, 1990c).

Nitzschia barbieri has only rarely been recorded since Peragallo (1921) and Hasle (1965a).
See discussion under Fragilariopsis rhombica.

187
 DIATOMS

Nitzschia bicapitata Cleve (1901a: 933, fig. 12)

ADDITIONAL REFERENCES: Hustedt (1958: 169, figs 176-190), Simonsen (1974: 50, pi. 35, figs 3-
15), Kaczmarska et al. (1986: 1859, fig. 10), Hasle & Medlin (1990b: 177, pi. 23.3, fig. 3), Hasle
& Syvertsen (1997: 330, pi. 74, tab. 77), Fryxell (2000: 47, figs 1-12)

Fig. 2.108a, b

DESCRIPTION: Cells solitary, lanceolate, with capitate apices; apical axis 8-27 um; transapical
axis 3.8-5.8 urn [see Hasle & Syvertsen (1997, tab. 77) and Fryxell (2000, fig. 2)].
Transapical striae straight in valve centre, but curved towards ends, 25-34 in 10 um, each a
uniseriate row of areolae; areolae 30-40 in 10 um. Fibulae 12-16 in 10 um. Raphe eccentric
(Fig. 2.108a). Central nodule present.

distribution: Coastal waters off Davis Station, East Antarctica; Southern Ocean, offshore
Davis Sea and offshore Wilkes Land; Weddell-Scotia Confluence (Garrison et al., 1987);
66°N-62°S (Hasle, 1976b).
Features that separate the different bicapitate Nitzschia species include cell dimensions and
the densities of fibulae, striae and areolae (Fryxell, 2000). Nitzschia bicapitata is further
characterised by having a row of small, rather inconspicuous pores (smaller than the areolae)
on both valve mantles.

Cells collected by us had slightly larger dimensions (apical axis 10.4-27 um; transapical axis
3.8-5.8 um; striae 30-34 in 10 um; areolae 30-35 in 10 um) than the emended description of
Fryxell (2000).

Nitzschia decipiens Hust. (Hustedt, 1958: 170, figs 174, 175)

SYNONYMS: Nitzschia acicularis var. lantarctica van Heurck (1909: 19, pi. 3, fig. 63)
Nitzschia dissipata var. antarctica Perag. (Peragallo, 1921: 65, pi. 3, fig. 14)

ADDITIONAL REFERENCE: Hasle & Medlin (1990b: 177, pi. 23.3, fig. 3)

description: Cells solitary, lanceolate in valve view, with rostrate ends; apical axis 30^0
um; transapical axis 2-2.7 um. Transapical striae straight, close-set, 44 in 10 um. Fibulae 7-9
in 10 um. Central nodule absent.
distribution: Type locality, 69°43'S 06°30'E (Hustedt, 1958); Antarctic (Hasle & Medlin,
1990b).

Morphologically similar to N. acicularis, but rarely reported from southern waters.

Nitzschia lecointei van Heurck (1909: 21, pi. 3, fig. 57)

SYNONYMS: Nitzschia acicularis var. interrupta Heiden & Kolbe (1928: 670)
Nitzschia lanceolata var. spicula Manguin (1960: 334, pi. 19, fig. 232)
Nitzschia lanceolata var. pygmaea f. australis Manguin (1960: 334, pi. 19, figs 230, 231)

Nitzschia acicularis f. angustior auct. non Miiller (1905), sensu Helmcke & Krieger (1954: 17, pi. 190)

ADDITIONAL references: Hustedt (1958: 172, pi. 13, figs 167, 168), Hasle (1964: 23, pi. 1,

fig. 1; pi. 4, fig. 5; pi. 5, fig. 2; pi. 11, figs 14, 15; pi. 12, figs 9-13), Hasle & Medlin (1990b: 177,
pi. 23.2, figs 1-5; pi. 23.3, figs 1, 2)

Fig. 2.109a-e

description: Cells solitary; median part rhomboid to lanceolate in valve view, with rostrate
ends; apical axis 15-112 jam; transapical axis 2.5-5 um. Transapical striae straight, close-
set, 51-55 in 10 um. Interstrial membranes perforated by 1 row of small poroids, c. 60 in 10

um. Fibulae 5-14 in 10 um. Central nodule present. See Hasle (1964) for a more detailed
description.

distribution: Near Davis Station, East Antarctica and Cape Evans; Ross Sea; type locality,
 DIATOMS

Figure 2.108. Nitzschia species, a, b, N. bicapitata. a, SEM Cell in valve view, b, LM Cell
in valve view, c, N. longissima, SEM Two mediocomtricta. d, SEM
solitary cells, d-f, N.
Solitary cell, oblique valve view with raphe (arrows), e, SEM Cell in broad girdle view, f,
SEM Partly collapsed cell in broad girdle view. Scale bars: a, b = 2 urn; c-f = 10 urn.

SO
 DIATOMS

Figure 2.109. Nitzschia lecointei. a. SEM Two cells, b. SEM Cell in oblique valve and broad
girdle view. c. SEM Single cell. d. SEM Fibulae and striae in middle part of cell. e. TEM
Fibulae and striae at cell end. Scale bars: a-c = 5 urn; d, e = 1 urn.

90
 DIATOMS

sea-ice, 65°15.5'S 64°30.0

,
W (van Heurck, 1909); Indian sector of the Antarctic (Kozlova,
1962); Terre Adelie (Frenguelli, 1960); Antarctic (Hasle & Medlin, 1990b); Weddell-Scotia
Confluence (Garrison et ah, 1987).

The most common habitat of N. lecointei appears to be the undersurface of ice

(tychopelagic). When liberated from this substratum, it will vegetate as a plankton form near
the ice-edge.

is confined to the zone nearest the ice

In the Pacific part of the Antarctic Ocean, N. lecointei
and was recorded sporadically from water samples. Originally, it was sampled "par la fusion
de glace de mer, colore en vert-brun fonce" (van Heurck, 1909). Heiden & Kolbe (1928)
reported N. acicularis var. interrupta from ice as well as in quantitative samples. Manguin
(1960) reported the smaller N. lanceolata var. pygmaea f. australis as being common in
bottom mud ("boue de profondeur") at Cape Margerie, and the larger var. spicula as very
rare in the same habitat. Frenguelli (1960) noted the species as rare in three samples from
Terre Adelie, while Kozlova (1962) documented small amounts in water samples from
stations close to Antarctica in the Indian Ocean sector.

Nitzschia longissima (Breb.) Ralfs, in Pritchard (1861: 783, pi. 4, fig. 23)
BASIONYM: Ceratoneis longissima Breb., in Kiitzing (1849: 891)
See VanLandingham (1978: 3072) for further synonymy.

ADDITIONAL REFERENCES: van Heurck (1880-85: 185, pi. 70, figs 1-4), Peragallo (1897-1908:
293, pi. 74, figs 16-20), Cupp (1943: 200, fig. 154), Proschkina-Lavrenko (1955b: 200, fig. 95),
Hasle (1964: 20, pi. 1, fig. 2; pi. 5, fig. 6; pi. 10, figs 5-7; pi. 11, figs 1-4), Hasle & Syvertsen
(1997:329, pi. 74)

Fig. 2.108c

DESCRIPTION: Cells solitary; median part narrowly lanceolate in valve view, with slightly
125-450 pm; transapical axis 6-7 urn. Transapical striae 52-60 in
rostrate ends; apical axis
10 pm (Hasle, 1964). Fibulae 6-14(?) in 10 um, narrow and arcuate, adjoining silicified ribs
of the same diameter which run parallel to the raphe fissure (see Hasle, 1964, pi. 11, fig. 2).
Central nodule present.

distribution: Southern Ocean, south of Australia; Indian Ocean sector of the Antarctic
(Kopczyhska et ah, 1986).

See notes under Cylindrotheca closterium.

Nitzschia medioconstricta Hust. (Hustedt, 1958: 174, figs 165, 166)

Fig. 2.108d-f

description: Cells solitary and tapered towards ends in apical view, distinctly constricted in
the middle and with truncate ends in girdle view; apical axis 55-86 pm; transapical axis 5-7
pm. Transapical striae straight, 26-30 in 10 pm. Fibulae 8-12 in 10 pm.
distribution: Plough I. and coastal waters off Davis Station, East Antarctica; in stomach
contents of Salpa fusiform is in the Antarctic, 69°43'S 06°30'E (Hustedt, 1958).

Hustedt (1958) suggested that N. medioconstricta is related to the 'W. hybrida group", which
includes N. hybrida Grunow, N. subhybrida Hust., N. hybridaeformis Hust. and
N. pseudohybrida Hust.

At first glance, N. medioconstricta can be confused with Auricula compacta, particularly in

girdle view. However, the latter is usually a much longer cell (79-134 pm), and it has fewer
fibulae (6-9 per 10 pm).

91
 DIATOMS

Nitzschia peragallii Hasle (1965b: 41, pi. 9, figs 11, 12; pi. 15, figs 1-5)
SYNONYM: Nitzschia barbieri var. latestriata Perag. (Peragallo, 1921: 66, pi. 3, fig. 21)

ADDITIONAL REFERENCE: Hasle & Medlin (1990c: 181, pi. 24.4, figs 11, 12; pi. 24.7, figs 1-5)

DESCRIPTION: Cells solitary or in short chains with cells attached by valve faces, elliptical to
lanceolate in valve view; apical axis 32-90 |im; transapical axis 7-8 um. Transapical striae
straight in the middle of the cell, curved near valve poles, 7-9 in 10 um. Interstrial
membranes with 2 rows of poroid areolae; areolae c. 16 in 10 um. Fibulae present in the
same density as striae. Pseudonodulus absent.

distribution: Type locality, Argentine Is., Antarctic Peninsula (Peragallo, 1921; Hasle,
1965b); Antarctic pack-ice, Bellingshausen Sea, BRATEGG (Hasle, 1965b).

This species has rarely been reported since its original description.

Nitzschia sicula (Castrac.) Hust. var. bicuneata (Grunow) Hasle (1960: 26, fig. 16c, d)

SYNONYMS: Raphoneisl (Diatoma?) bicuneata Grunow, in Cleve & Moller (1879: nos 208-210)
Pseudonitzschia sicula var. bicuneata (Grunow) Perag., in Peragallo & Peragallo (1897-1908: 299, pi. 72, fig. 26)

Nematoplata bicuneata (Grunow) Kuntze (1898: 416)

See Hasle (1960, 1964) for a more detailed description and additional synonymy.

additional REFERENCES: Hasle (1964: 38, figs 11, 12; pi. 5, fig. 8; pi. 13, fig. 14; pi. 16, fig. 1),

Hasle & Syvertsen (1997: 327, pi. 74)

Fig. 2.110a, b

DESCRIPTION: Cells solitary, lanceolate, with somewhat protracted ends in valve view; apical
axis 32-39 um; transapical axis 6 um. Transapical striae coarse, 8-12 in 10 um. Interstrial
membranes with 2-5 rows of small poroids close to the interstriae. Fibulae 8-12 in 10 um.
Raphe eccentric; central raphe endings terminating in a circular depression. Central nodule
present.

distribution: Southern Ocean, 57°35'S 105°46'E, offshore Wilkes Land; type locality,
Rovingo, Adriatic Sea (Cleve & Moller, 1879); Antarctic BRATEGG stations (Hasle, 1964).

Because their morphology is poorly known, Hasle & Syvertsen (1997) place four varieties of
N. sicula within a group "Incertae sedis {Nitzschia)". Simonsen (1974) noted that this species
is highly variable in general (valve) outline. He also observed transitions between
var. bicuneata and var. migrans (Cleve) Hasle in his "Meteor" samples.

Nitzschia stellata Manguin (1957: 132, pi. 7, fig. 47a-e)

ADDITIONAL references: Manguin (1960: 335, pi. 19, figs 234-236; pi. 31, figs 379, 380),
McConville et al. (1985: 137, as N. frigida); Medlin & Hasle (1990a: 160, pi. 21.1, fig. 4; pi.

21.3, fig. 3)

Fig. 2.1 1 la-e

description: Cells solitary or attached end-to-end and forming straight, zig-zag or stellate
colonies of 3-12 cells, linear to lanceolate in valve view, tapering towards the poles; apical
axis 60-134 um; transapical axis 6.5-10.5 um. Transapical striae c. 34—40 in 10 um,
scarcely visible by LM. Fibulae 4-8 in 10 um. Central nodule present.

distribution: Type locality, Terre Adelie (Manguin, 1957); sea-ice near Davis Station, East
Antarctica (Archer etal., 1996b); Antarctic ice (Medlin & Hasle, 1990b); coastal waters off

Syowa Station, East Antarctica (Ishikawa et al., 2001).

It is likely that many Antarctic reports of N . frigida Grunow, a Northern Hemisphere species,
should be referred to N. stellata.

Nitzschia taenia W.Sm. (Smith, 1853: 43, pi. 15, fig. 123)

SYNONYM: Nitzschia taeniiformis Simonsen (1978: 525)

92
 DIATOMS

Figure 2.110. Nitzschia sicula var. bicuneata. a, SEM Solitary cell, valve view with
eccentric raphe (arrows), b, SEM Detail of poroid interstrial areas and central endings of
raphe (arrow). Scale bars: a, b = 2 um.

ADDITIONAL references: Hustedt (1958: 180, fig. 195), Medlin & Hasle (1990a: 160, pi. 21.1,
fig. 5; pi. 21.4, fig. 2, as N. taeniiformis)

description: [after Hustedt (1958)] Cells solitary or attached by valve faces to form ribbon-
like colones, linear and tapering towards the poles in valve view, rectangular in girdle view;
apical axis 110-150 um; transapical axis 5 urn. Transapical striae 24 in 10 fim. Fibulae 6-10
in 10 um. Central nodule present.

distribution: Antarctic ice (Medlin & Hasle, 1990a).

We did not collect this diatom in our investigations, and it has rarely been reported from
Antarctic waters.

Pseudonitzschia antarctica Manguin (1960: 329, pi. 19, figs 218, 219a, b)

description: [after Manguin (I960)] Cells solitary, linear, weakly sigmoidal, with rounded
apices (valve view); apical axis c. 100 um; transapical axis 3-4 um. Fibulae 14 in 10 um;
transapical striae not visible by LM. Central nodule present.

distribution: Terre Adelie (Manguin, 1960).

This rare species was not collected by us. Manguin (1960) noted the features separating
P. antarctica from the Arctic diatom Nitzschia delicatissima Cleve [= P. delicatissima
(Cleve) Heiden], the former being larger and weakly sigmoidal.

Pseudonitzschia heimii Manguin (1957: 131, pi. 6, fig. 43a-c)

SYNONYMS: Nitzschia pacifica Cupp (1943: 204, fig. 157), Hustedt ( 1958: 177, pi. 13, figs 193, 194)

Nitzschia heimii (Manguin) Hasle (1965a: 21, pi. 1, figs 6, 7; pi. 10, figs 9-13; pi. 11, figs 1-7), Hasle &
Medlin (1990a: 169, pi. 22.2, figs 9-13)

193
 DIATOMS

Figure 2.111. Nitzschia LM Vegetative cells in stellate pattern, b, SEM Vegetative

stellata. a,
cells in stellate pattern, c, Vegetative cell, valve view, d, SEM Vegetative cell, girdle
SEM
view, e, SEM Girdle view with fibulae (arrows) protruding from raphe. Scale bars: a-c = 20
um; d = 10 urn; e = 1 urn.

194
 DIATOMS

Figure 2.112. Pseudonitzschia heimii. a, LM Pair of cells, b, SEM Single cell in oblique
view, c, SEM Apex of cell with eccentric raphe (arrow), d, SEM Internal surface of valve

showing fibulae (arrows) and finely poroid striae, e, SEM Detail of transapical striae. Scale
bars: a, b = 10 urn; c-e =1p.m.

195
 DIATOMS

Figure 2.113. Pseudonitzschia species, a-e, P. lineola. a, LM Chain of cells, b, SEM Chain
of cells, c, SEM Overlapping ends of 2 cells, d, SEM Overlapping ends of 2 cells: fibulae
(arrows) visible through broken part of cell, e, SEM Overlapping ends of 2 cells, f, g, P.
prolongatoides. f, TEM Solitary cell, g, TEM Cell apex; fibulae (arrows) visible. Scale bars:
a, b = 10 urn; c-e = 2 \im; f = 5 urn; g = 1 um.

196
 DIATOMS

ADDITIONAL REFERENCES: Manguin (1960: 332, pi. 19, figs 223, 224), Hasle & Syvertsen (1997: 315, pi. 70,
tab. 74)

Fig. 2.1 12a-e

DESCRIPTION: Cells solitary or in long rigid chains with c. a quarter of cell ends overlapping,
linear to lanceolate and attenuated towards the poles in valve view, with one margin straight;
cell ends somewhat sigmoidal and obliquely truncated at the poles in girdle view; apical axis
67-120 um; transapical axis 4-6.5 um. Transapical striae broad, usually strongly silicified,
19-26 in 10 um. Interstrial membranes perforated by or 2 rows of minute poroids, c. 50-80
1

in 10 um; poroids in 2 or 3 rows on valve mantle and valvocopulae. Raphe keel eccentric;
fibulae distinct, 1 1-16 in 10 um. Central nodule present.

DISTRIBUTION: Southern Ocean, south of Australia; Heard I.; type locality, coastal Terre
Adelie (Manguin, 1957); Antarctic and sub-Antarctic (Hasle & Medlin, 1990a); North
Atlantic (Hasle, 1965a); KERFIX, 50°40'S 68°25'E, Southern Ocean (Kopczyhska et al,
1998); Weddell-Scotia Confluence (Garrison et al, 1987).

Specimens from the most northerly sub- Antarctic stations (Hasle, 1965a) had 70-80 small
poroids in 10 |am and rather broad intervals between the transapical rows of poroids.

Pseudonitzschia lineola (Cleve) Hasle (1965a: 29, pi. 12, figs 15, 16)

BASIONYM: Nit:schia lineola Cleve (1897: 300, fig. 10), Hasle & Medlin (1990a: 170, pi. 22.3, figs 15-21)

SYNONYMS: Nitzschia barkleyi Hust. (Hustedt, 1952: 293, figs 13, 14)

Pseudonitzschia barkleyi (Hust.) Manguin (1960: 330)

ADDITIONAL REFERENCES: Manguin (1954: 23, Heard I.), Hasle (1974: 319), Hasle & Syvertsen
(1997: 317, pi. 71, tab. 75)

Fig. 2.113a-e

description: Cells solitary or in long rigid chains, overlapping c. one-sixth of cell length,
linear to lanceolate in girdle view, linear and attenuated towards the poles in apical view,
with slightly sigmoidal ends; apical axis 56-112 um; transapical axis 1.8-2.7 um.
Transapical striae 22-28 in 10 um. Interstrial membranes perforated by 1 (or 2) rows of
minute poroids. Keel eccentric; fibulae distinct, 11-16 in 10 um. Central nodule present.

DISTRIBUTION: Coastal waters off Davis Station and Polar Aviation I., East Antarctica; type
60°03'N 05°51'W (Cleve, 1897); Antarctic, sub- Antarctic, Arctic (Hasle & Medlin,
locality,
1990a); KERFIX, 50°40'S 68°25'E, Southern Ocean (Kopczynska et al, 1998); Weddell-
Scotia Confluence (Garrison et al., 1987).

Pseudonitzschia prolongatoides (Hasle) Hasle (1993: 319)

BASIONYM: Nitzschia prolongatoides Hasle (1965a: 25, pi. 1, fig. 8; pi. 2, figs 11-13; pi. 12, figs 7, 8; pi. 13,
figs 1, 2; text figs 1, 2)

SYNONYM: Nitzschia prolongata Manguin (1957: 132, pi. 7, fig. 46), auct. non Hustedt (1938: 489, pi. 41, fig. 28)

ADDITIONAL REFERENCES: Hasle (1964: pi. 5, fig. 10), Hasle & Medlin (1990a: 170, pi. 22.3, figs
7, 8), Hasle & Syvertsen (1997: 317, pi. 73, tab. 75)

Fig. 2.1 13f, g

description: Cells solitary or in stellate colonies, linear to rostrate in valve view, often with
slightly inflated apices and expanded in middle area; apical axis 20-70 um; transapical axis
0.5-2.5 um. Transapical striae 30-35 in 10 p.m. Interstrial membranes with 2 or 3 rows of
minute poroids. Keel eccentric; fibulae 15-18 in 10 um. Central nodule present. [See Hasle
(1965a) for a full description.]

DISTRIBUTION: Sea-ice near Davis Station, East Antarctica (Archer et al, 1996); Terre Adelie
(Manguin, 1957); Antarctic (Hasle & Medlin, 1990a); Weddell-Scotia Confluence (Garrison
etal, 1987).

197
 DIATOMS

Pseudonitzschia prolongatoides is similar in appearance to P. subcurvata, but in valve view

the latter has one straight margin and one curved, whereas prolongatoides is inflated evenly
in the centre of the cell.

Pseudonitzschia subcurvata (Hasle) G.A.Fryxell (1993: 319)

BASIONYM: Nitzschia subcurvata Hasle (1964: 28, pi. 1, fig. 3; pi. 12, figs 14-16; pi. 13, figs 1-6)
SYNONYM: Nitzschia closteriodes Hasle (1956: 616) pro parte (see Hasle, 1964: 28)

ADDITIONAL REFERENCES: Hasle (1974: 319), Hasle & Medlin (1990a: 177, pi. 23.2, figs 6-8),
Fryxell et al. (1991: 243), Hasle & Syvertsen (1997: 323, pi. 73, tab. 75)

Fig. 2.114a-f

description: Cells solitary or in chains, then with a short overlapping of valve ends, one
margin straight in valve view, the other convex, thus inflated in middle part of valve; apical
axis 47-113 urn; transapical axis 1.5-2.5 urn. Transapical striae Fine, 44-49 in 10 urn, not
visible by LM. Keel eccentric; fibulae irregularly spaced, 12-18 in 10 urn. Central nodule
absent. [See Hasle (1964) for a full description.]

distribution: Inshore waters off Davis Station, East Antarctica; Southern Ocean, offshore
Wilhelm II Land at 60°50'S 93°36'E, and offshore Prydz Bay at 67°01'S 77°43'E; Antarctic
(Hasle & Medlin, 1990a); Lutzow-Holm Bay (Tanimura et al., 1990); Weddell-Scotia
Confluence (Garrison et al., 1987).

Auxospore formation was described in detail and illustrated by Fryxell et al. (1991).

Pseudonitzschia turgidula (Hust.) Hasle (1993: 320)

BASIONYM: Nitzschia turgidula Hust. (Hustedt, 1958: 182, pi. 13, figs 172, 173)

ADDITIONAL REFERENCES: Hasle (1965a: 24, pi. 1, fig. 11; pi. 2, fig. 3; pi. 11, figs 8-13; pi. 12,
figs 1-6; 1974: 320), Hasle & Medlin (1990a: 169, tab. 22.1, pi. 22.3, figs 1-6), Hasle &
Syvertsen (1997: 319, pi. 71, tab. 75)

Fig. 2.1 15a-c

DESCRIPTION: Cells solitary or chain-forming, overlapping c. one-sixth of cell length, linear in

apical view, but gradually expanded towards the middle section; apices rounded. In girdle
view margins slightly convex and ends truncate (Fig. 2.115c); apical axis 30-80 urn;
transapical axis 2.5-3.5 urn. Transapical striae 23-28 in 10 urn. Interstrial membranes with 2
rows of minute poroids (Fig. 2.115b). Raphe keel eccentric; fibulae 13-18 in 10 urn. Central
nodule present.

distribution: Southern Ocean, offshore Wilhelm II Land at 60°50'S 93°36'E; type

localities, South Atlantic 46°32'S 0°02'W and 51°29'S 0°08'W (Hustedt, 1958); Pacific
sector of the Southern Ocean, 53-62°S, BRATEGG (Hasle, 1965a).

The closely related P. turgiduloides is larger and has more coarsely structured fibulae and
striae (see below).

Pseudonitzschia turgiduloides (Hasle) Hasle (1993: 320)

BASIONYM: Nitzschia turgiduloides Hasle (1965a: 28, pi. 12, figs 9-14; pi. 13, figs 3-6), Hasle & Medlin
(1990a: 170, pi. 22.3, figs 9-14)

SYNONYM: Pseudonitzschia barkleyi var. obtusa Manguin (1960: 332, pi. 19, fig. 222)

ADDITIONAL REFERENCE: Hasle (1974: 320), Hasle & Syvertsen (1997: 323, tab. 75)

Fig. 2.1 15d-f

DESCRIPTION: Cells chain-forming, linear in valve view, with rounded apices, slightly
expanded in girdle view; apical axis 63-126 um; transapical axis 1.2-2.7 urn. Transapical

198
 DIATOMS

Figure 2.114. Pseudonitzschia subcurvata. a, LM Single cell, b, SEM Single cell, c, TEM
Single cell, d, TEM Fibulae (arrows) and striae at cell apex, e, TEM Fibulae (arrows) and
striae near middle of cell, f, SEM Striae (arrows) near middle of cell. Scale bars: a-c = 10
urn; d-f =1 um.

199
 DIATOMS

Figure 2.115. Pseudonitzschia species, a-c. P. turgidula. a, SEM Solitary cell in valve view.
b, SEM Detail of valve surface, c, SEM Solitary cell in girdle view, d-f, P. turgiduloides. d,
SEM Short chain of 2 cells, e, SEM Overlapping cell ends. f. TEM Overlapping cell ends.
Scale bars: a, c, d = 10 um; b, e, f =
1 urn.

200
 DIATOMS

striae broad, 17-21 in 10 urn. Interstrial membranes with 1 or 2 rows of minute poroids.
Raphe keel eccentric; fibulae 10-13 in 10 urn. Central nodule present. [See Hasle (1965a) for
a full description.]

distribution: Coastal waters off Davis Station, East Antarctica; Antarctic (Hasle & Medlin,
1990a); Weddell-Scotia Confluence (Garrison et al., 1987); coastal waters off Syowa Station,
East Antarctica (Ishikawa et al., 2001).

Pseudonitzschia turgiduloides is similar to P. delicatissima (Cleve) Heiden from the Arctic

and the somewhat more fine-structured P. turgidula.

201
 3. DINOFLAGELLATES*
Andrew McMinn & Fiona J. Scott

[Order Suessiales by Paul G. Thomson]

Dinoflagellates are small (usually less than 100 urn), unicellular, motile cells most of which
exhibit a characteristic 'whirling' motion in water. They possess two flagella, one located
around the circumference of the cell in a groove called the girdle, and the other orientated to
the posterior of the cell. This well-defined group of organisms has comparatively few
representatives in Antarctic waters in contrast to temperate and tropical seas. Both thecate
(those with a wall composed of plates) and athecate forms are found. The abundance of
dinoflagellates in Antarctic waters can be substantial and seasonal (Lessard & Rivkin, 1986;
Bjornsen & Kuparinen, 1991; McKenzie & Cox, 1991). Dinoflagellates can be heterotrophic,
autotrophic or mixotrophic, and the distinction is readily made by using fluorescence
microscopy to observe the autofluorescence of the cells. Thus, autotrophic organisms contain
chloroplasts that fluoresce red under blue light, whereas heterotrophs do not. However,
caution should be exercised so as not to misinterpret chloroplasts ingested with food as being
being part of the dinoflagellate.

The taxonomy followed here is that adopted by Steidinger & Tangen (1997) who noted that
morphology and cytology are important criteria for the identification of dinoflagellate
species, but they also confirmed the usefulness of biochemical and genetic markers. In cases
where more recent examination of material has resulted in revised taxonomy (e.g. Polarella),
the systems of later authors have been followed.

The following dinoflagellates, reported only rarely from Antarctic waters, are not treated in
detail here:

Blepharocysta paulsenii J.Schiller (see Dodge & Priddle, 1987)

Ceratium horridum (Cleve) Gran (see Balech, 1968)

C. kofoidii Jorg. (see Hada, 1970; Okamoto & Takahashi, 1984)
Okamoto et al., 1986)
C. peters ii Nielsen (see

Dinophysis simplex Bohm (see Okamoto et al., 1986)

Preperidinium granulosum (Balech) Elbr. (see Dodge & Priddle, 1987)
Protoperidinium acanthophorum (Balech) Balech (see McKenzie & Cox, 1991)
P. joergensenii (Balech) Balech (see McKenzie & Cox, 1991)
P. macrapiticum (Balech) Balech (see Dodge & Priddle, 1987; Andreoli et al., 1995)
P. manginii (Balech) Balech (see Watanabe, 1982; Dodge & Priddle, 1987)
P. pallidum (Ostenf.) Balech (see Watanabe, 1982; Dodge & Priddle, 1987)
P. parcum (Balech) Balech (see Watanabe, 1982; Dodge & Priddle, 1987)
P. parvicollum (Balech) Balech (see Watanabe, 1982; Dodge & Priddle, 1987)
P. pseudoantarcticum (Balech) Balech (see Watanabe, 1982; Dodge & Priddle, 1987)

P. pyriforme (Paulsen) Balech (see McKenzie & Cox, 1991)

P. pyrum (Balech) Balech (see Watanabe, 1982; Dodge & Priddle, 1987)

P. radius Balech (see Priddle, 1985; Andreoli et al., 1995)

P. raphanum (Balech) Balech (see Watanabe, 1982; Dodge & Priddle, 1987)

P. saltans (Meunier) Balech (see Balech, 1976; Watanabe, 1982)

P. sphaeroideum (Mangin) Balech (see McKenzie & Cox, 1991)
P. turbinatum (Mangin) Balech (see Watanabe, 1982; Dodge & Priddle, 1987)
P. variegatum (N.Peters) Balech (see Watanabe, 1982; Dodge & Priddle, 1987)

Class Dinophyceae or Subphylum Dinoflagellata.

202
 DINOFLAGELLATES

Order Prorocentrales

Family Prorocentraceae

Mesoporos perforatus (Gran) Lillick (1937: 497)

BASIONYM: Exuviaella perforata Gran (1915: 99, fig. 7)

SYNONYMS: Porella bisimpressa J.Schiller (1928: 54, fig. 12), nom. Meg.
Porella perforata J.Schiller (1928: 55, fig. 12), nom. Meg.
Porella adriatica J.Schiller (1928: 56, fig. 1
1 ), nom. Meg.
Porella globulus J.Schiller (1928: 56, fig. 13), nom. Meg.
Porella asymmetrica J.Schiller (1933: 27, fig. 29), nom. Meg.
Porotheca perforata (Gran) Silva (1960: 23)

ADDITIONAL REFERENCES: Dodge (1982: 24, text fig. 2A, B; pi. la, b); Steidinger & Tangen
(1997:419)

Fig. 3.1a, b

description: Cells round in valve view, flattened, 14-17 urn long, 15-21 urn diameter. Epitheca
and hypotheca each with a central pore. Girdle ribbed, 2-5 urn wide, with 2 flagellar pores.
Theca densely covered with small papillae; area around the central pore smooth. Periphery of
each valve with a row of pores. Chloroplasts 2, one each in the epitheca and hypotheca.

distribution: Southern Ocean, 43-63°S, south of Australia.

Prorocentrum
Three species, antarcticum, P. balticum and P. compressum, have been reported from
P.
Antarctic waters. Prorocentrum is characterised by small, simple, rounded to elliptical cells

and a cell wall of only 2 thecal plates. Dodge & Bibby (1973), Sournia (1986) and other
authors placed Exuviaella in synonymy with Prorocentrum.

Figure 3.1. Mesoporos perforatus. a, SEM Oblique girdle view, b, SEM Valve view of theca
and central pore. Scale bars: a, b = 10 urn.

203
 DINOFLAGELLATES

Figure 3.2. Prorocentrum species, a, P. antarcticum, TEM Valve view, b, c, P. balticum,

SEM Valve view (b) and oblique apical view (c). d, P. compressum, TEM Valve view. Scale
bars: a-d = 5 urn.

204
 DINOFLAGELLATES

Prorocentrum antarcticum (Hada) Balech (1975: 15)

BASIONYM: Exuviaella antarclica Hada (1970: 10, fig. 3)

SYNONYM: Exuviaella sp., Balech & El-Sayed (1965: 112, pi. 3, fig. 55)

ADDITIONAL REFERENCE: Balech (1976: 27, fig. 16)

Fig. 3.2a

description: Cells ovate, flattened, c. 15-22 um long, c. 15-25 um diameter. Theca smooth,
with 2 short anterior projections; cell surface sparsely covered with small pores. Chloroplasts
present.

distribution: Southern Ocean, coastal regions to north of the Polar Front (Balech, 1975);
KERFIX, 50°40'S 68°25'E, Southern Ocean (Kopczyhska et ai, 1998); sea-ice near Davis
Station, East Antarctica (Archer et ai, 1996a, as Exuviaella antarclica); Admiralty Bay,
King George I., Antarctic Peninsula (Brandini, 1993).

Prorocentrum antarcticum is distinguished from P. compressum (see below) by its smaller

size and fewer pores. It has only been observed as a minor component of Southern Ocean
dinoflagellate communities. Estrada & Delgado (1990) reported P. cf. antarcticum from the
Weddell Sea.

Dodge (1975: 107, fig. 1A) considered Exuviaella antarctica Hada to be synonymous with
P.aporum (J.Schiller) J.D.Dodge, but the latter is not a widely accepted combination.

Prorocentrum balticum (Lohmann) A.R.Loebl. (Loeblich, 1970: 906)

BASIONYM: Exuviaella baltica Lohmann (1908: 265, pi. 17, fig. la, b)

SYNONYMS: Prorocentrum pomoideum Bursa (1959: figs 108-1 1 1)

Exuviaella aequatoralis Hasle (1960: 29, fig. 18a, b)

additional reference: Dodge (1982: 35, text fig. 2M, N; pi. le)

Fig. 3.2b, c

description: Cells round in valve view, 9-22 um long, 14-21 um diameter, not or barely
compressed in side view; older cells with a large intercalary band. Periflagellar area with a
triangular flagellar pore and a smaller accessory pore.

distribution: Southern Ocean, offshore Wilkes Land, 57°35'S 105°46'E and Southern
Ocean south of Australia.
The specimens collected by us were slightly larger than previously recorded. Cells 9-10 um
long were reported by Dodge (1982) and less than 20 um by Steidinger & Tangen (1997).

Prorocentrum compressum (Bailey) Abe ex J.D.Dodge (1975: 1 10, text figs 2F, 4H, I;

pi. 4E, F)

BASIONYM: Pyxidicula compressa Bailey (1850: 40, pi. 2, figs 13, 14)

SYNONYMS: Exuviaella compressa Ostenf. (Ostenfeld, 1899: 59)

Exuviaella oblongata J.Schiller (1928: 50, fig. 6a-c)

Prorocentrum bidens J.Schiller (1928: 61, fig. 21)
Prorocentrum lebourae J.Schiller (1928: 62, fig. 23)
Exuviaella lenticulata Matzen. (Matzenauer, 1933: 438, fig. 1)

Exuviaella elongata Rampi (1951 : pi. 1, fig. 9)

additional reference: Dodge (1982: 31, fig. 21)

Fig. 3. 2d

description: Cells broadly ovate in valve view, 30-50 um long, c. 25 um diameter,

compressed (sometimes only slightly) in side view. Surface covered with small pores.

205
 )

DINOFLAGELLATES

distribution: Southern Ocean, south of Australia, 63°14'S 120°19'E; widely distributed in

the Southern Ocean, but not found south of the Polar Front (Steidinger & Tangen, 1997);
sub-Antarctic (Balech, 1976).

Order Dinophysales

Family Amphisoleniaceae

Dinophysis
Dinophysisis a distinctive genus with up to ten representatives in the Southern Ocean. While

diversity lower than in many other marine environments, occasionally species of Dinophysis
is

have been observed to dominate Southern Ocean dinoflagellate floras (Dodge & Priddle,
1987).

The genus is characterised by a small rounded to flat epitheca and a much larger, rounded to

ellipticalhypotheca. The cingulum is well defined and bordered by prominent lists. Lists are
also associated with the sulcus which, because of the lenticular shape of the cell, are usually
located on the side.

Many authors consider Phalacroma to be synonymous with Dinophysis (e.g. Tai &
Skogsberg, 1934; Abe, 1967). The genera overlap in many aspects of their morphology, but
can be separated the distinctive, funnel-shaped, anterior cingular list of Dinophysis. Other
features, such as the development and direction of cingular lists in combination with the
height and shape of the epitheca, also help to distinguish the genera. Members of the genus
Dinophysis are specialised predators, and Hansen (1991) observed D. rotundata Clap. &
J.Lachm. [= Phalacroma rotundatum (see below)] and D. hastata Stein ingesting the
prostomatid ciliate Tiarina fusus (Clap. & J.Lachm.) Bergh.

Dinophysis antarctica Balech (1958a: 82, pi. 2, figs 14-25), as antarcticum

ADDITIONAL REFERENCES: Balech (1976: 29, fig. 18), Andreoli et al. (1995: 473, fig. 18)

Fig. 3.3a, b

description: Cells rounded to elliptical in side view, 28-65 um long, 20-55 urn diameter.
Epitheca very low and generally flat. Sulcal lists smooth; left sulcal list always narrow at the
lower septal ray (R3); right sulcal list reduced. Cell surface covered with areolae and scattered
pores; additional apical or antapical projections (excrescences) absent. Chloroplasts present.

distribution: Southern Ocean, south of Australia; Terra Nova Bay, Ross Sea (Andreoli et
al.,1995); Southern Ocean, near South Georgia (Dodge & Priddle, 1987); Bellingshausen
Sea (Balech, 1973); Weddell Sea, Antarctic Peninsula (Balech, 1973, 1975).

Dinophysis antarctica can be distinguished from most other Antarctic species by its flat
epitheca and lack of excrescences. The otherwise similar D. punctata is much smaller, while
D. meteori is almost rectangular in shape. This species has been observed only infrequently
in eastern Antarctica, some specimens being somewhat smaller than previously reported.

Dinophysis contracta (Kofoid & Skogsb.) Balech (1973: 8, pi. 1, figs 6-17)
BASIONYM: Phalacroma contractum Kofoid & Skogsb. (Kofoid & Skogsberg, 1928: 83, fig. 3, 1

SYNONYMS: Phalacroma ruudii Braarud (1935: 1 12, fig. 32)

Prodinophysis contracta (Kofoid & Skogsb.) Balech (1944: 429)

Prodinophysis ruudii (Braarud) A.R.Loebl. (Loeblich, 1965: 17)

ADDITIONAL REFERENCES: Balech (1976: 35, fig. 23), Dodge (1982: 48, fig. 4H; pi. 2b)

Fig. 3.3c, d

206
 DINOFLAGELLATES

Figure 3.3. Dinophysis species, a, b, D. antarctica, SEM Cell (a) and detail of theca surface
(b). c, d, D. contracta, SEM Solitary cells. Scale bars: a, c, d = 10 urn; b = 1 urn.

207
 DINOFLAGELLATES

Figure 3.4. Dinophysis species, a, D. meteori, SEM. b, D. operculoides, SEM. c, D. ovum.

SEM. d, D. punctata, SEM. Scale bars: a-d = 10 (im.

208
 DINOFLAGELLATES

description: [see Dodge (1982) for a detailed description] Cells small, rounded to
ellipsoidal, 20-30 urn long, 17-25 um diameter. Epitheca large and conical, occupying up to
Cingulum 5 urn wide, deeply incised, smooth, with rows of
one-third the length of the cell.
marginal pores; cingular lists greatly reduced. Sulcal lists narrow, inconspicuous by LM.
Cell surface densely covered with pores; apical and antapical excrescences absent.
Chloroplasts present.

distribution: Southern Ocean, south of Australia; Southern Ocean, near South Georgia
(Dodge & Priddle, 1987); Falkland Is. and the south-eastern Atlantic (Balech, 1988).
The large, conical epitheca distinguishes this from most other Dinophysis species.
Dinophysis tenuivelata also has a prominent epitheca, but it lacks a deeply incised cingulum.
This is the most common Dinophysis species in coastal waters of eastern Antarctica;
however, it never comprises more than 10 percent of a dinoflagellate community.

Dinophysis cornuta (N.Peters) Balech (1967: 82)

BASIONYM: Phalacroma cornuta N.Peters (1928: 20, fig. 1)

SYNONYM: Phalacroma spinata N.Peters (1928: 22, fig. 2)

ADDITIONAL REFERENCE: Balech (1976: 33, fig. 21)

description: [after Balech (1976)] Cells medium-sized with an irregular rounded to

triangular outline, 45-64 um long, 40-50 urn diameter. Epitheca large, rounded.
Excrescences well developed and irregular at apex and antapex. Left sulcal list wide, flared
at the base (at R3). The 2 upper sulcal rays (Rl and R2) close together. Cingulum moderately
indented. Cell surface covered with pores. Chloroplasts present.

distribution: Southern Ocean, near South Georgia (Dodge & Priddle, 1987); confined to
south of the Polar Front (Balech, 1976).

This uncommon but distinctive species was not seen by us. Balech (1976) recognised
subsp. cornuta and subsp. inerme Balech (in Balech & El-Sayed, 1965: 1 15, pi. 1, figs 12-18),
based on the development of excresences. The latter lacks apical excrescences and has only
inconspicuous antapical excrescences. The tubular outgrowths usually present in the dorsal
and antapical sagittal regions of D. norvegica Clap. & J.Lachm. from Northern Hemisphere
waters are similar to those of D. cornuta (Taylor & Pollingher, 1987). Dinophysis cornuta is
relatively easy to identify because of its irregular outline caused by pronounced apical and
antapical excrescences and by its extended left sulcal list.

Dinophysis meteori B6hm (1933: 16, fig. 2)

additional reference: Balech (1976: 31, fig. 19)

Fig. 3.4a

description: Cells medium-sized, rectangular, 45-55 um long, 45-50 um diameter.

Epitheca very short, flat. Cingulum deeply incised. Cell surface densely covered with pores;
apical and antapical excrescences absent. Chloroplasts present.

distribution: Southern Ocean, south of Australia; Ross Sea (Watanabe, 1982); Bransfield
Strait (Priddle, 1985); Southern Ocean, near South Georgia (Dodge & Priddle, 1987).

This species can be distinguished from D. antarctica by its more rectangular outline and
deeply incised cingulum. It is most commonly found north of the Polar Front.

Dinophysis operculoides (Schutt) Balech (1967b: 83)

BASIONYM: Phalacroma operculoides Schutt (1895, pi. 2)

ADDITIONAL REFERENCE: Balech (1988: 49, pi. 10, figs 1 1-15)

Fig. 3.4b

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 DINOFLAGELLATES

description: Cells medium-sized, rounded, 40-62 urn long, c. 40 um diameter. Epitheca

very reduced, flat to slightly rounded. Cingulum 7 um wide, not incised; anterior cingular
list wider (4 um) than posterior list, both lists smooth and projected anteriorly. Sulcal lists

smooth; left sulcal list extending for half cell length, c. 8 um wide, with 3 visible rays; right
sulcal list c. half length of and narrower than the left list. Cell surface comparatively smooth,
with scattered pores. Chloroplasts present.

distribution: Southern Ocean, south of Australia, 44-60°S; South Atlantic (Balech, 1988);
South Georgia (Dodge & Priddle, 1987).

Dinophysis ovum Schutt (1895: pi. 10, fig. 6)

SYNONYM: Dinophysis brevisulcus Tai & Skogsb. (Tai & Skogsberg, 1934: 430, fig. 3a-k)

ADDITIONAL REFERENCES: Abe (1967: 50, fig. 10a-p), Dodge (1982: 53, fig. 3J)

Fig. 3.4c

description: Cells medium-sized, ovoid, 40-58 um long, 30-45 um diameter. Epitheca

obscured by anterior cingular list. Hypotheca rounded, covered with deep areolae, each with
a pore. Cingulum 4 um wide, the areolae with pores, incised on the dorsal side only; anterior
cingular list forming a funnel over the epitheca, wider than posterior list, both lists smooth
and projected anteriorly. Sulcal lists smooth; left sulcal list extending for half cell length,
c. 8 um wide, with 3 visible rays; right sulcal list c. one-third length of the left list (to the

second ray) and narrower. Chloroplasts present.

distribution: Southern Ocean, south of Australia, 43-63°S; Antarctic waters, south of the
Polar Front (Hara & Tanoue, 1985).

Dinophysis punctata Jorg. (Jorgensen, 1923: 23, fig. 28)

ADDITIONAL REFERENCES: Lebour (1925: 81, pi. 12, fig. 5), Schiller (1933: 118, fig. Ill), Balech
(1976: 29, fig. 17), Dodge (1982: 54, fig. 3 I), Balech (1988: 41, pi. 5, figs 19-21)

Fig. 3.4d

description: Cells small, rounded to ellipsoidal, 30-40 um long, 25-35 um diameter.

Epitheca very reduced, flat to slightly rounded. Cingulum not incised. Cell surface covered
with scattered pores. Chloroplasts present.

distribution: South of the Polar Front; Southern Ocean, near South Georgia (Dodge &
Priddle, 1987); Bellingshausen Sea and Weddell Sea, around the Antarctic Peninsula
(Balech, 1976).

A small, poorly understood species without many well-defined distinguishing features. It is

most similar to D. antarctica from which it can be separated by its smaller size.

Dinophysis tenuivelata Balech (1973: 10, pi. 1, figs 18-20; pi. 2, figs 21-26)

additional REFERENCE: Balech (1976: 36, fig. 24)

Fig. 3.5a, b

description: Cells medium-sized, rounded to ellipsoidal, 35-45 um long, 35-45 um

diameter. Epitheca rounded, well developed, comprising up to 20% of the cell length.
Cingulum not incised. Sulcal lists narrow, inconspicuous by LM. Cell surface densely
covered with pores; apical and antapical excrescences absent.

distribution: Infrequent throughout the Southern Ocean; Antarctic (Balech, 1976).

Dinophysis tenuivelata can be distinguished from D. contracta by its larger size, smaller and
rounded (as opposed to conical) epitheca and non-incised cingulum. There can be difficulty
in separating D. tenuivelata from Phalacroma rotundatum (q.v.), but the latter is usually
more ellipsoidal and has a somewhat flattened epitheca.

210
 DINOFLAGELLATES

Figure 3.5. Dinophysis and Heteroschisma species, a, b, D. tenuivelata, SEM. c. D. tuberc-

ulata, SEM. d, H. subantarctica, SEM. Scale bars: a-d = 10 ixm.

211
 DINOFLAGELLATES

Figure 3.6. Unidentified Gymnodinium species, SEM. Scale bars: a-d = 5 um.

212
 DINOFLAGELLATES

Dinophysis tuberculata Mangin (1926: 72, fig. 15)

ADDITIONAL reference: Balech (1976: 31, fig. 20)

Fig. 3.5c

description: Cells large, irregular to ellipsoidal, 55-82 urn long, 48-59 urn diameter.
Epitheca very reduced and flat. Cingulum incised. Sulcal lists and excrescences broad and
irregular (R3 is particularly well developed). Cell surface covered with scattered pores.

distribution: Southern Ocean, 46-63°S, Antarctic; seldom north of the Polar Front (Balech,
1976).

Heteroschisma subantarctica Balech (1971: 48, pi. 4, figs 39-48)

additional references: Balech (1976: 36, fig. 25; 1988: 38, pi. 4, figs 1 1-13)

Fig. 3.5d

description: Cells rounded, c. 20 urn long, c. 17 um diameter. Epitheca clearly visible

above the cingular list. Hypotheca c. 4 times the length of the epitheca. A characteristic large
pore visible on the epitheca above the sulcus. Cingulum 4-5 um wide, deeply incised;
anterior and posterior cingular lists smooth, comparatively narrow and projected laterally.
Left sulcal list c. three-quarters length of hypotheca, with 3 clearly visible rays. Surface
areolate, with large scattered pores on both thecae.

distribution: Southern Ocean, 44-62°S, south of Australia; Antarctic (Balech, 1976).

Phalacroma rotundatum (Clap. & J.Lachm.) Kofoid & J.R.Michener (1911: 290)
BASIONYM: Dinophysis rolundata Clap. & J.Lachm. (Claparede & Lachmann, 1859: 6, pi. 20, fig. 16)

SYNONYMS: Prodinophysis rotundata (Clap. & J.Lachm.) Balech (1944: 429, fig. 2)

Dinophysis whittingae Balech (1971: figs 154-167)

ADDITIONAL REFERENCES: Dodge (1982: 55, fig. 41; pi. 1 if), Balech (1988: 46, pi. 9, figs 1-4,
as D. rotundata), Steidinger & Tangen (1997: 439, pi. 14)

DESCRIPTION: [after Steidinger &

Tangen (1997)] Cells small, rounded to ellipsoidal, 36-56
pm long, 35-40 pm diameter. Epitheca rounded, sometimes flattened at the apex. Cingulum
5-8 pm wide, slightly incised; anterior cingular list a little wider than posterior list; both
lists project laterally with rays that appear to be extensions of the surface ornamentation of
the lists. Left sulcal list slightly wider and longer than right sulcal list; left sulcal list with
3 visible rays, with the right list extending to the third ray. Cell surface (thecae and
cingulum) areolate; each areola with a pore. Chloroplasts absent.

DISTRIBUTION: Southern Ocean 43-63°S, south of Australia; Southern Ocean, near South
Georgia (Dodge & Priddle, 1987); Ross Sea (Watanabe, 1982).

We did not collect Phalacroma rotundatum, a species that is known to have a similar
morphology to Dinophysis contracta (q.v.). Many authors consider Phalacroma to be
synonymous with Dinophysis.

Order Gymnodiniales

Family Gymnodiniaceae

Amphidinium hadai Balech (1976: 19, fig. 10)

SYNONYM: Amphidinium acutissimum auct. non Schiller (1933), sensu Hada (1970: 12, fig. 5)

ADDITIONAL REFERENCE: Andreoli et al. (1995: 473)

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 DINOFLAGELLATES

DESCRIPTION: Cells fusiform, 26-49 urn long, 3-13 urn wide. Epitheca short, ±conical.
Hypotheca c. 4 times larger than epitheca, acuminate. Sulcus short and inconspicuous.
Chloroplasts numerous, large and ellipsoidal.

distribution: Terra Nova Bay, Ross Sea (Andreoli et al., 1995); KERFIX, 50°40'S
68°25'E, Southern Ocean (Kopczyiiska et al., 1998); Antarctic pack-ice (Garrison, 1991b).

A small but distinctive species, A. hadai is the only common Amphidinium species in
Antarctic waters.

Gymnodinium
These are unarmoured, solitary or chain-forming cells. The cingulum is usually equatorial or
premedian, with or without descending (left) displacement; if displaced, then less than 20%
of cell length (cf. Gyrodinium) (Steidinger & Tangen, 1997). See Larsen (1994, 1996) for a
discussion of the separation of Gymnodinium and Gyrodinium.

Individual species are differentiated by their size, shape and contours, chain formation, the
occurrence and shape of the apical groove, the presence of striae, ribs or furrows, cingulum
premedian, median or postmedian, the displacement of the cingulum, the extent to which the
sulcus extends onto the epitheca, the presence of chloroplasts, lateral compression of the
body and colour (Steidinger & Tangen, 1997). Identification is difficult, and we include here
a series of images representing the range of morphologies seen (Figs 3.6 and 3.7).

Gymnodinium baccatum Balech, in Balech & El-Sayed (1965: 1 14, pi. 1, figs 4, 5)

ADDITIONAL REFERENCE: Balech (1976: 15, fig. 4)

distribution: Weddell Sea (Balech & El-Sayed, 1965); coastal waters near Syowa Station,
East Antarctica (Hada, 1970).

Gymnodinium frigidum Balech, in Balech & El-Sayed (1965: 1 12, pi. 1, figs 6, 7)

ADDITIONAL REFERENCE: Balech (1976: 15, fig. 3)

distribution: Southern Ocean, near South Georgia (Dodge & Priddle, 1987).

Gymnodinium modestum Balech (1976: 15, fig. 5)

SYNONYM: Gymnodinium baccatum auct. non Balech (1965), sensu Hada (1970: 13, fig. 8)

distribution: South of the Polar Front (Balech, 1976); coastal waters near Syowa Station,
East Antarctica (Hada, 1970).

Gymnodinium soyai Hada (1970: 13, fig. 9)

ADDITIONAL REFERENCE: Balech (1976: 18, fig. 7)

distribution: South of the Polar Front (Balech, 1976); coastal waters near Syowa Station,
East Antarctica (Hada, 1970).

Gyrodinium
Unarmoured sometimes compressed. Cingulum displaced by at least 20-35% of the
cells,
cell length (cf. Gymnodinium)
in a descending left spiral. Individual species are
differentiated by size, shape and contours, the presence and shape of the apical groove, the
presence of striae, ribs or furrows, total displacement of the cingulum, torsion of the sulcus,
the presence of chloroplasts, lateral compression of the body and colour (Steidinger &
Tangen, 1997). See Larsen (1994, 1996) for a discussion of the separation of Gymnodinium
and Gyrodinium.

214
 DINOFLAGELLATES

Figure 3.7. Unidentified Gymnodinium species, SEM. Scale bars: a-d = 5 u.m.

215
 DINOFLAGELLATES

Figure 3.8. Gyrodinium species, a, b, G. glaciale, SEM Ventral views, c, G. lachryma, SEM
Dorsal view, d, e, G. rhabdomante, LM Cell in two focal planes. Scale bars: a-c = 5 urn; d, e
= 10 urn.

216
 b

DINOFLAGELLATES

Gyrodinium glaciale Hada (1970: 16, fig. 13a, b)

ADDITIONAL REFERENCE: Balech (1976: 23, fig. 15)

Fig. 3.8a,

description: Cells small, biconical, 22-42 um long, 12-15 um wide {fide Balech, 1976),
fusiform and with pointed apices. Girdle offset by approximately one-third of the cell length.

distribution: Terra Nova Bay (Andreoli et al., 1995); Antarctic (Balech, 1976).

Balech (1976) illustrated this species with small, irregular chloroplasts.

Gyrodinium lachryma (Meunier) Kofoid & Swezy (1921: 314, fig. EE6)
BASIONYM: Spirodinium lachryma Meunier (1910: 63, pi. 14, figs 21, 22)

ADDITIONAL REFERENCES: Balech (1976: 21, fig. 12), Dodge (1982: 102, fig. 12C), Andreoli et
al. (1995: 473), Steidinger & Tangen (1997: 453, pi. 20)

Fig. 3.8c

DESCRIPTION: Cells large, 60-135 um long, 28-50 \im wide. Epitheca and hypotheca ±equal
in size; epitheca conical to tear-drop-shaped; hypotheca rounded. Girdle broad, incised and
offset by c. one-quarter of the cell length. Cell wall thick.

distribution: Coastal areas of Prydz Bay, East Antarctica, including saline lakes; Terra Nova
Bay (Andreoli et al., 1995); Weddell Sea (Estrada & Delgado, 1990, as G. cf. lachryma);
Antarctic pack-ice (Garrison, 1991b); Arctic, Antarctic and eastern U.S.A. (Dodge, 1982).

Balech (1976) reported the presence of small, irregular chloroplasts.

Gyrodinium rhabdomante Balech (1973: 7, pi. 1, figs 4, 5)

ADDITIONAL REFERENCE: Balech (1976: 21, fig. 13)

Fig. 3.8d, e

description: Cells large, distinctive, biconical, 150-170 um long, 60-88 um wide. Epitheca
and hypotheca pointed, ±equal in size, with prominent longitudinal ribs.

distribution: Coastal waters of Prydz Bay, East Antarctica; Bellingshausen Sea (Balech,
1976).

A rarely reported species. Balech (1976) noted the presence of 2 long, slender, conspicuous,
refringent rods that are fused together at the apex.

Katodinium sp.

ADDITIONAL REFERENCE: Steidinger & Tangen (1997: 453)

Fig. 3.9a, b

description: Cells unarmoured, fusiform, 55-60 um long, 18-20 um diameter. Theca with
c. 8 longitudinal ridges; epitheca tapering towards a pointed apex, approximately three-
quarters of the cell length. Cingulum descending. Chloroplasts absent.

distribution: Southern Ocean, south of Australia.

A small number of specimens were seen in the present study, but unambiguous identification
was not possible.

Torodinium sp.

Fig. 3.9c

217
 DINOFLAGELLATES

description: Cells unarmoured, elongate, cigar-shaped, 85 um long, 18 um diameter. Theca

ornamented with c. 20 longitudinal ridges; epitheca c. two-thirds of the cell length, tapering
towards a rounded apex. Cingulum descending. Chloroplasts not seen.

distribution: Southern Ocean, south of Australia.

We found a single specimen but could not identify it with certainty. "Torodinium sp." was
recorded for the Weddell Sea by Estrada & Delgado (1990).

Family Polykrikaceae

Polykrikos schwartzii Butschli (1873: 657)

ADDITIONAL REFERENCES: Sournia ( 1986: 60), Steidinger & Tangen (1997: 457, pi. 21)

Fig. 3.9d, e

description: Cells large, unarmoured, ovoid, 90-130 um long, 50-65 um diameter;

a pseudocolonial cell of 8 closely appressed zooids, each with its own cingulum and
transverse flagellum, but all sharing a common continuous sulcus. Chloroplasts absent.

distribution: Southern Ocean, South of Australia.

Order Suessiales

Family Suessiaceae

Polarella glacialis Montresor, Procaccani & Stoecker (1999: 187, figs 1-13, 20-22)

ADDITIONAL REFERENCES: Buck et al. (1992), Stoecker et al. (1997: 591, fig. 5; 1998), Thomson
et al. (2004: 867)

Fig. 3.10a-d

description: [modified from Montresor et al. (1999)] Vegetative cells: 10-15 um long,
6-9 um wide, with a rounded episome and a conical hyposome. Cell surface smooth, with a
few small scattered protuberances; amphiesmal plates concealed by outer membrane.
Cingulum deep, left-handed, displaced by c. cingular width. Sulcus shallow, not extending
1

into episome. Insertion point of flagella obscured by protrusion of the episome.

Chloroplast(?s) present; it is unclear whether there is a single multilobed chloroplast or many
chloroplasts converging towards the central part of the cell.

Cysts: Prolate-spherical, 12-17 um long, 8-15 um wide, with numerous recurved spines.
Paracingulum equatorial, lacking spines and slightly displaced.

distribution: Drygalski Glacier, Ross Sea; in fast-ice near Casey, Davis and Mawson
Stations and Larsemann Hills, East Antarctica (Thomson et al., unpublished data); abundant
in brine pockets and channels of upper fast-ice (Stoecker et al., 1997, 1998; Thomson &
McMinn, unpublished data); McMurdo Sound (Stoecker et al., 1997, 1998); pack-ice in the
Indian sector of the Southern Ocean, 62°56.87'S 92°58.00'E (Thomson et al., 2004);
Weddell and Scotia Seas (Garrison & Buck, 1989b; Buck et al., 1992); bipolar and
circumpolar in the Antarctic (Montresor et al., 2003).

Larger cysts (to 25 um diameter), morphologically indistinguishable from P. glacialis, have

been reported from sea-ice near Davis Station and McMurdo Sound. Cysts with surface pores
have been documented (e.g. Stoecker et al., 1992, fig. 9A), and Montresor et al. (1999)
suggested that these cysts might be closely related species or, possibly, phenotypes of
P. glacialis.

218
 DINOFLAGELLATES

Figure Katodinium, Torodinium and Polykrikos species, a, Katodinium sp., LM Solitary

3.9.
cell, Katodinium sp., LM Dividing cell, c, Torodinium sp., SEM Solitary cell, d, e,
b,
Polykrikos schwartzii, LM. Scale bars: a-e = 10 jxm.

219
 DINOFLAGELLATES

Figure 3.10. Polarella glacialis. a, LM Cysts, b, SEM Aggregation of cysts from sea-ice. c,

SEM Cyst, d, SEM Motile cell. Scale bars: a-d = 5 urn.

220
 1

DINOFLAGELLATES

Order Gonyaulacales

Family Ceratiaceae

Ceratium
A distinctive, armoured genus with long, apical and antapical horns, a prominent cingulum,
clear tabulation and longitudinal ribs (Jorgensen, 1911). Tabulation of Ceratium is typically
4', 7", 6c, 12s, 6'", lp, 1"". sometimes dorsally compressed and
The central body is

chloroplasts are present, although some species have food vacuoles due to phagocytic
activity. Only a few taxa have been collected from Antarctic waters, the most prominent
being the C. pentagonumlC. lineatum complex and the C. extensumlC. fusus complex.

Ceratium extensum (Gourret) Cleve (1900b: 15)/Ceratium fusus (Ehrenb.) Dujard.

(1841: 378) complex

ADDITIONAL REFERENCES: Dodge (1982: 231), Steidinger & Tangen (1997: 472)

Fig. 3.1 la-d

DESCRIPTION: Cells large, fusiform, 250 urn to more than 1 long, 15-25 urn diameter; mm
right antapical horn small or absent; left hypothecal horn straight. Cingulum c. 10 urn wide,
only slightly excavated, ornamented with pores; cingular lists very narrow. Cell surface
generally smooth, with scattered pores; small spines are sometimes present.

distribution: Southern Ocean, 43-62°S, south of Australia; south of 70°S (Balech, 1968);
cosmopolitan in cold-temperate to tropical waters (Steidinger & Tangen, 1997).

There is considerable morphological overlap between Ceratium extensum and C. fusus. Both
are large fusiform dinoflagellates with fully developed apical and left antapical horns but
reduced right antapical horns. The difference between the two is based on the relative length
of the hypotheca; in C. extensum it is distinctly longer than the epitheca. However, we found
this character to be unreliable.

Okamoto & Takahashi (1984) did not find this complex south of 50°S, and they reported it to
be rare at water temperatures below 7°C.

Ceratium pentagonum Gourret (1883: 1 1)/Ceratium lineatum (Ehrenb.) Cleve

(1899: 36) complex

ADDITIONAL REFERENCES: Dodge (1982: 230, fig. 28F; pi. 8d), Steidinger & Tangen (1997: 477,
pi. 26)

Fig. 3.12a-c

description: Cells dorsiventrally flattened, with a broadly pentagonal outline, 120-200 urn
long, 50-80 urn diameter. Epitheca convex adjacent to cingulum, but tapering to a long apical
horn c. twice body length. Hypotheca supporting 2 stout antapical horns c. equal to body
length, the left horn a little longer than the right. Epitheca and hypotheca approximately equal
in size, separated by an entrenched cingulum bordered by prominent cingular ridges.
Cingulum slightly helical. Sulcus prominent, occupying more than half of ventral surface.
Surface ornamented with a prominent series of longitudinal ridges extending to tips of horns.
Tabulation typical of Ceratium, 4', 7", 6c, 12s, 6'", lp, 1"". Chloroplasts present.

distribution: Adjacent to and immediately south of the Polar Front; Southern Ocean, near
South Georgia (Dodge & Priddle, 1987); Southern Indian Ocean, 46-62°S, 75°E (Okamoto et
al, 1986).

There is considerable morphological overlap between Ceratium pentagonum and C lineatum

and, as a consequence, we are treating them here as a single complex. Both have ±pentagonal

221
 DINOFLAGELLATES

outlines, but C. pentagonum usually has a longer apical horn, although we found this
character to be quite variable.

Ceratium pentagonum can be distinguished from C. fusus by its shape. Although typically an
oceanic, warm-temperate to tropical species (Steidinger & Tangen 1997), C. pentagonum is

sometimes the most abundant dinoflagellate around the Polar Front (Balech, 1976) and South
Georgia (Dodge & Priddle, 1987).

Ceratium pentagonum subsp. grande Mangin was recorded for the Antarctic by Balech
(1976:90, fig. 66).

Ceratium tripos (O.F.Mull.) Nitzsch (1817: 4)

BASIONYM: Cercaria tripos O.F.Mull. (Miiller, 1776: 206)

ADDITIONAL REFERENCES: Lebour (1925: 148, pi. 32, 33), Schiller (1937: 382, figs 384, 385),
Sournia (1967: 406, fig. 27), Dodge (1982: 234, fig. 30A-D), Steidinger & Tangen (1997: 478, pi.

26)

Fig. 3.12d

description: Cells 70-90 urn long overall, 50 urn diameter, with a sharp transition from the
triangular epitheca to the60-90 urn long apical horn. Hypotheca rounded to flattened at the
antapex, tapering longitudinally into curved antapical horns; antapical horns 50-70 urn long,
the left usually longer than the right. Cingulum sometimes reduced to a line, sometimes
slightly excavated or with narrow lists. Sulcus a large aperture on the ventral side, extending
into the epitheca. Surface ornamentation variable, smooth or reticulate, and with scattered
pores, spines or horns.

distribution: Southern Ocean, 44-6 1°S, south of Australia.

Family Goniodomataceae

Alexandrium tamarense (Lebour) Balech (1995: 38, pi. 6, figs 1-40; pi. 7, figs 1-9)

BASIONYM: Gonyaulax tamarensis Lebour (1925: 95, pi. 14, fig. 1)

synonyms: Pyrodinium phoemts Wolosz. & W.Conrad (Woloszyriska & Conrad, 1939: 3, figs 1-12)
Gonyaulax excavata (Braarud) Balech (1971 : 28, pi. 7, figs 1 19-124)
Gonyaulax globosa (Braarud) Balech (1971: 29)
Gonyaulax trygvii Parke, in Parke & Dodge (1976: 550)
Gessnerium tamarensis (Lebour) A.R.Loebl. & L.A.Loebl. (Loeblich & Loeblich, 1979: 44)

Protogonyaulax excavata (Braarud) F.J. R.Taylor (1979: 51)

additional REFERENCES: Dodge (1982: 216, fig. 25A-C), Steidinger & Tangen (1997: 500, pi. 36)

Fig. 3.13a-d

description: Cells rounded to ovoid, 20-44 (am long, 16-36 urn diameter. Thecae thin and
delicate; epitheca broadly conical, rounded at the apical pore; hypotheca rounded with
convex sides. Cingulum equatorial c. 5 urn wide, displaced by 0.5-1.5 cingular widths.
Cingular lists narrow, smooth; anterior list projected posteriorly; posterior list projected
anteriorly. Chloroplasts present.

distribution: Southern Ocean, 50-62°S, south of Australia; coastal waters off Davis
Station, East Antarctica.

Cysts of this species have been reported from north-eastern Japan by Ishikawa & Taniguchi
(2000).

222
 DINOFLAGELLATES

Figure 3.11. Ceratium externum! C. fusus complex, a, C. extensum, SEM Single cell, b, C.
externum, SEM Transverse sulcus, c, C. fusus (specimen from southern Australia), SEM
ventral view, d, C. fusus (specimen from southern Australia), SEM dorsal view. Scale bars:
a, c, d = 50 urn; b = 5 urn.

223
 DINOFLAGELLATES

Figure 3.12. Ceratium species, a-c. C. pentagomnnlC. lineatum complex, a. SEM Ventral
view, b, c, SEM Dorsal views, d. C. tripos. SEM Dorsal view. Scale bars: a-d = 10 urn.

224
 DINOFLAGELLATES

Figure 3.13. Alexandrium tamarense. a, SEM Oblique antapical view, b, SEM Ventral view,
with flagella.c, SEM Oblique antapical view, d, SEM Ventral view. Scale bars: a-d = 5 um.

225
 DINOFLAGELLATES

Family Gonyaulacaceae

Gonyaulax digitale (Pouchet) Kofoid ( 1 9 1 1 : 2 1 4, pi. 9, figs 1-5)

BASIONYM: Protoperidinium digitale Pouchet (1883: 443, pi. 18, fig. 14)

ADDITIONAL REFERENCE: Dodge (1982: 208, fig. 26A; pi. 6c, d)

Fig. 3.14a

description: Cells rhomboidal, 50-75 pm long, 35-50 pm diameter. Thecae thick, with
large deep pores scattered over the surface; epitheca triangular with ±straight sides grading
into a short apical horn; hypotheca rounded, with slightly convex sides, with 2 antapical
spines, the left spine longer than the right. Cingulum c. 5 pm wide, displaced by 2-2.5
cingular widths; cingular lists absent or greatly reduced, narrow, projected laterally. Sulcus
deep, extending from apex to antapex; sulcal lists narrow. Chloroplasts present.

distribution: Southern Ocean, 50-62°S, south of Australia; Southern Ocean, near South
Georgia (Dodge & Priddle, 1987); Ross Sea (Watanabe, 1982).

Gonyaulax digitale typically has epithecal shoulders and a moderate apical horn.

Gonyaulax scrippsae Kofoid (191 1: 228, pi. 13, figs 26, 27)

ADDITIONAL REFERENCES: Dodge (1982: 214, fig. 26A, H), Steidinger & Tangen (1997: 507, pi. 42)
description: Cells rounded to ovoid, 29-50 pm long, 27-40 pm diameter. Thecae marked
with raised longitudinal ridges; epitheca with convex sides and a short apical horn with an
apical pore; hypotheca rounded, lacking antapical spines. Cingulum displaced by 2-3
cingular widths, with strong ridges; lists greatly reduced or absent. Sulcus narrow, deeply
indented, extending from apex to antapex; sulcal lists ±absent. Chloroplasts present.

distribution: Cysts have been recorded from the Southern Ocean; Southern Ocean, near
South Georgia (Dodge & Priddle, 1987); Ross Sea (Watanabe, 1982).

Figure 3.14. Gonyaulax digitale and Scrippsiella troichoidea. a, G. digitale, SEM Ventral
view, b,S. troichoidea, SEM Cyst. Scale bars: a = 10; b = 2 pm.

226
 DINOFLAGELLATES

Cysts of this species have the palaeontological name Spiniferites bulloideus (Deflandre &
Cookson) Sarjeant (Wall & Dale, 1968). These are ovoid, 32-42 pm long, 28-39 urn
diameter, with bifid and trifid sutural spines c. 16 pm long.

Gonyaulax striata Mangin (1926: 74, fig. 1611)

ADDITIONAL REFERENCE: Dodge (1989: 289, figs 2J, 22)

Fig. 3.15a-d

description: Cells almost pentagonal, 20-30 pm long, 16-25 pm diameter. Thecal surfaces
with strong longitudinal ridges or striae interconnected by scattered pores; epitheca with
±straight sides tapering to a small apical horn with an apical pore, broad at the base;
hypotheca rounded, without antapical spines. Cingulum c. 4 pm wide, displaced by 1.5
cingular widths; cingular lists absent or greatly reduced, narrow and ribbed. Sulcus narrow,
extending from apex to antapex; sulcal lists ±absent except at antapex where they extend
posteriorly. Chloroplasts present.

distribution: Southern Ocean, 44-55°S, south of Australia; Southern Ocean, near South
Georgia (Dodge & Priddle, 1987); Ross Sea (Watanabe, 1982).

Family Oxytoxaceae

Oxytoxum criophilum Balech, in Balech & El-Sayed (1965: 121, pi. 4, figs 51-54)

description: [modified from Balech & El Sayed (1965)] Cells 56-61 pm long (epitheca
11.5-14 pm), 33-36 pm maximum width. Epitheca low, triangular, with somewhat convex
sides, abruptly tapering to a short spine. Hypotheca reticulate, with longitudinal ribs and a
short antapical spine. Cingulum excavated, slightly overhanging; cingular lists absent. Sulcus
very short; sulcal lists absent.

distribution: Weddell Sea (Balech & El-Sayed, 1965).

Rarely reported and not seen by us. Dodge & Saunders (1985) partially revised the genus
Oxytoxum based on SEM studies.

Order Peridinales

Family Calciodinellaceae

Scrippsiella troichoidea (J. Stein) A.R.Loebl. (Loeblich, 1976: 25)

BASIONYM: Glenodinium troichoideum J. Stein (1883: pi. 3, figs 27-29)
SYNONYMS: Glenodinium acuminatum Jorg. (Jorgensen 1899: 32)

Peridinium troichoideum (J. Stein) Lemmerm. (Lemmermann, 1910: 336)

Scrippsiella faeroense Dickensh. & Ed. R. Cox (Dickensheets & Cox, 1971: 139, figs 1-6)

ADDITIONAL REFERENCES: Wall et al. (1970: 151), Dodge (1982: 163, fig. 18Q-S; pi. 3d),
Steidinger & Tangen (1997: 527, pi. 6, 47)

Fig. 3.14b

description: Cells pyriform, 16-36 pm long, 20-23 pm diameter. Thecae with scattered
poroids; epitheca conical, with a short convex apical process and collar; hypotheca rounded.
Cingulum broad, indented, with narrow cingular lists. Chloroplasts present.
Cysts calcareous, spherical, 7-8 pm diameter, densely covered with short pointed or
terminally flattened spines.

distribution: Southern Ocean, 64°S, south of Australia; widely distributed in middle and
low latitudes of the Northern Hemisphere (Wall et al, 1970).

227
 DINOFLAGELLATES

Family Kolkwitziellaceae

Diplopsalis lenticula Bergh (1882: 244, figs 60-62)

SYNONYMS: Glenodinium lenticula (Bergh) Pouchet (1883: 44, pi. 20, 21)

Dissodinium lenticulum (Bergh) A.R.Loebl. (Loeblich 1970: 905)

additional REFERENCES: Dodge (1982: 154, fig. 18I-K; pi. 3b-d), Steidinger & Tangen (1997:
529, pi. 48)

description: [after Dodge (1982)] Cells subspherical to lenticular, 23-48 um long, 32-68 um
diameter; surface smooth with scattered pores. Cingulum equatorial; cingular lists
comparatively broad, projecting laterally. Left sulcal list prominent, curved to the right.
Tabulation la, 6", 4c, 5s, 5'", 1""; plate orthoform. Chloroplasts absent.

distribution: Southern Ocean, near South Georgia (Dodge & Priddle, 1987).

Rarely reported and not seen by us.

Preperidinium meunierii (Pavill.) Elbr. (Elbrachter, 1993: 176)

BASIONYM: Peridinium meunierii Pavill. (Pavillard, 1912: 177)

SYNONYMS: Diplopeltopsis lenticula f. minor Paulsen (1907: 9, fig. 9)

Diplopeltopsis minor (Paulsen) Pavill. (Pavillard, 1913: 7)

Zygabikodinium lenticulatum Loebl. & A.R.Loebl. (Loeblich & Loeblich, 1970: 541)

ADDITIONAL references: Balech (1976: 39, fig. 26), Steidinger & Tangen (1997: 530, pi. 48)

Fig. 3.16a-d

description: Cells medium-sized, lenticular, 20-30 um long, 40-50 um diameter, with an

almost circular cross section. Small apical horn present but no antapical horns. Apical pore
well developed, bordered by 2 shoulders. Cingulum not helical, not incised, bordered by
crenulate lists. Left sulcal list prominent. Cell surface covered with fine pores. Tabulation
lpa, 3', 2a, 7", 3c, 6s?, 5'", 1'; 1' plate orthoform, small and rhombic in shape; 2a plate
exceptionally large, occupying most of the apical dorsal surface.

DISTRIBUTION: Southern Ocean, near South Georgia (Dodge & Priddle, 1987; Bunt & Wood.
1968); Ross Sea and Syowa Station, East Antarctica (Watanabe, 1982); cosmopolitan
(Balech, 1968).

The very large antapical plate, which covers most of the hypotheca, is highly distinctive.
Cysts of this species have the palaeontological name Dubridinium caperatum F.Reid (Wall &
Dale, 1968).

Preperidinium perlatum (Balech) Elbr. (Elbrachter, 1993: 177)

BASIONYM: Diplopeltopsis perlata Balech (1971: 81, pi. 13, figs 198-207)

SYNONYM: Zygabikodinium perlatum (Balech) Elbr. (Elbrachter, 1975: 58-64)

ADDITIONAL REFERENCE: Balech (1976: 40, fig. 27)

Fig. 3.17a, b

description: Cells small, subspherical, 15-20 um long, 15-20 um diameter. Apical horn very
short; antapical horns absent. Cingulum broad, non-ascending, bordered by narrow unornam-
ented ridges. Sulcal lists inconspicuous. Cell surface densely covered with short (to 0.5 um
long) conical spines of 2 types; one set broader than the other (distinguishable only by SEM).
Tabulation lpa, 3', 2a, 7", 3c, 6s?, 5'", 2""; preapical plate within the apical pore small but
distinct; 2a plate very large; la plate pentaform.

DISTRIBUTION: Prydz Bay, East Antarctica; Southern Ocean, near South Georgia (Dodge &
Priddle, 1987); Ross Sea (Watanabe, 1982); Lutzow-Holm Bay and Terre Adelie (Hada,
1970); south-western Atlantic (Balech, 1968).

228
 DINOFLAGELLATES

Figure 3.15. Gonyaulax striata, a, SEM Ventral view, b, SEM Dorsal view, c, SEM Apical
view, d, SEM Antapical view. Scale bars: a-d = 5 urn.

229
 DINOFLAGELLATES

Figure 3.16. Preperidinium meunierii. a, SEM Ventral view, b, SEM Antapical view. c. d,

SEM Apical views. Scale bars: a-d = 10 urn.

230
 DINOFLAGELLATES

The surface ornamentation and distinctive tabulation are diagnostic. The presence of
2 antapical plates help to separate it from much larger, lenticular P. meunierii.

Family Podolampaceae

Blepharocysta splendor-maris (Ehrenb.) Ehrenb. (Ehrenberg, 1873: 4)

BASIONYM: Peridinium splendor-maris Ehrenb. (Ehrenberg, 1860: 791)

ADDITIONAL REFERENCES: Dodge (1982: 254, fig. 33H), Balech (1988: 125, pi. 53, figs 16-19),
Steidinger & Tangen (1997: 533, pi. 7, 49)

Fig. 3.17c

DESCRIPTION: Cells ovoid to ellipsoidal, somewhat laterally compressed, 31-70 um long, 35-
65 um diameter. Epitheca and hypotheca difficult to distinguish due to absence of a
cingulum; thecal plates smooth, with scattered pores. Sulcus extending from equatorial line
to apical pore and to antapex; sulcal lists narrow, but expanding to c. 8 um long distally to
form false antapical spines. Tabulation 3', la, 5", 3c, 4 or 5s, 4 or 5'", 1"".

distribution: Southern Ocean, 56-63°S, south of Australia; Atlantic (Dodge, 1982); South
Atlantic (Balech, 1988).

Podolampas antarctica Balech, in Balech & El-Sayed (1965: 121, pi. 4, figs 56-64)

ADDITIONAL REFERENCE: Balech (1976: 92, fig. 68)

Fig. 3.17d

DESCRIPTION: Cells top-shaped, 88-100 um long (including spines), 16-20 um diameter.

Epitheca tapered, ending in an apical pore and a short apical spine, covered with scattered
pores. Hypotheca smooth, with several rows of densely packed pores at antapical margin,
rounded, with 3 conspicuous antapical spines, the longest c. 20 um; spines interconnected by
membranes. Cingulum scarcely visible. Tabulation 3', la, 5", 3c, 4-6s, 5'", 1"", la, 5.

distribution: Southern Ocean, 43-56°S, south of Australia; type locality, Weddell Sea
(Balech & El-Sayed, 1965); Antarctic (Balech, 1976); South Georgia (Dodge Priddle, 1987).&

Family Protoperidiniaceae

Protoperidinium
Protoperidinium is by far the most diverse dinoflagellate genus in Antarctic waters. Approx-
imately 40 species have been recorded, most of which are endemic. While a number of taxa,
e.g. P. antarcticum, are readily identifiable by their gross morphology, it is usually necessary
to obtain good SEM images to determine the tabulation pattern essential for accurate
identification. For most species, identification is based on the shape and configuration of the
first apical plate (1') and the second intercalary plate (2a). Table 3.1 summarises the plate

configuration of Antarctic Protoperidinium species.

Balech (1974) recognised three subgenera, Minusculum, Archaeoperidinium and Protoperid-

inium. Subgenus Minusculum, with only a single Antarctic species, P. defectum, is
characterised by having 6 precingular plates of which 6" is especially large. Only 2 anterior,
intercalary plates are present.

Subgenus Archaeoperidinium, represented by two Antarctic species, has 7 precingular plates

but only 2 anterior, intercalary plates.

Most Antarctic species which is characterised by

are referrable to subgenus Protoperidinium
7 precingular plates and 3 anterior, intercalary plates. Primarytaxonomic differentiation is
based on the shape of the first apical plate, which can be surrounded by 4 (orthoform),
5 (metaform) or 6 (paraform) other plates (Fig. 3.18). The next most significant attribute is

231
 DINOFLAGELLATES

Figure 3.17. Preperidinium, Blepharocysta and Podolampas species, a, b. Preperidinium

perlatum. a, SEM Ventral view, b, SEM Dorsal view, c, Blepharocysta splendor-mavis, SEM
Ventral view, d, Podolampas antarctica, SEM Dorsal view. Scale bars: a-d = 10 \im.

232
 DINOFLAGELLATES

the shape of the second anterior, intercalary plate which, once again, can be in contact with
4 (quadra), 5 (penta) or 6 (hexa) surrounding plates (Fig. 3.18). For accurate identification, it
is usually essential to determine the form of these two plates, this requiring both a dorsal and

ventral view or an apical view. The most common plate configuration among the Antarctic
species is metaform quadra (Table 3.1).

Quadra

Figure 3.18. Types of plate configuration in Antarctic species of Protoperidinium

subg. Protoperidinium.

Protoperidinium adeliense (Balech) Balech (1974: 65)

BASIONYM: Peridinium adeliense Balech (1958b: 396, pi. 4, figs 101, 102)

additional reference: Balech (1976: 83, fig. 63)

description: Cells pyriform, 42-68 pm long (excluding horns), 47-82 um overall, 36-56 urn
diameter. Thecae smooth, with small scattered spines and pores. Epitheca with concave
sides, tapering toan apical horn. Hypotheca rounded, with 2 close-set divergent antapical
spines; membranes present on spines. Cingulum c. 5 um wide, non-incised, displaced by half
a cingular width; cingular lists broad, ribbed only at the list-theca interface. Sulcus barely
extending into epitheca, but extending and broadening to antapex; sulcal lists well
developed. Tabulation lpa, 4', 3a, 7", 6c, 5'", lp, 1"". Plate configuration: paraform quadra.

distribution: Type locality, Terre Adelie (Balech, 1958b); Southern Ocean, near South
Georgia (Dodge & Priddle, 1987); Ross Sea (Andreoli et ai, 1995).

Protoperidinium affine (Balech) Balech (1974: 65)

BASIONYM: Peridinium affine Balech (1958b: pi. 4, figs 91-100)

ADDITIONAL REFERENCES: Balech (1976: 88, fig. 65), Andreoli et at. (1995: 473, fig. 20)

Fig. 3.19a

233
 DINOFLAGELLATES

Figure 3.19. Protoperidinium species, a, P. affine, SEM Apical view, b, P. applanatum, SEM
Ventral view, c, d. P. antarcticum, SEM Ventral and apical views. Scale bars: a-d = 10 |jm.

234
 DINOFLAGELLATES

description: Cells almost pentagonal, 44-62 pm long (excluding horns), 50-71 urn long
overall, 47-59 urn diameter. Thecae smooth, with scattered pores. Epitheca slightly convex,
tapering to a short apical horn. Hypotheca rounded; antapex flattened, with 2 divergent
antapical spines; membranes present on spines. Cingulum non-incised, displaced by
1 cingular width; cingular lists wide, ribbed. Sulcus barely extending into epitheca, but
extending and broadening to antapex; sulcal lists well developed. Tabulation lpa, 4', 3a, 7",
6c, 5'", lp, 1"" (2a penta, large). Plate configuration: paraform penta.

distribution: Southern Ocean, 46-56°S, south of Australia; type locality, Terre Adelie
(Balech, 1958b); Southern Ocean, near South Georgia (Dodge & Priddle, 1987); Ross Sea
(Watanabe, 1982; Andreoli et al, 1995).

Table 3.1. Plate configuration of Antarctic Protoperidinium subg. Protoperidinium.

Orthoform Metaform Paraform

Quadra P. anlarcticum P. applanation P. adeliense

p. latistriatum P. bellulum P. areolatum

P. charcoti P. defectum
P. concavum
P. cf. crassipes

P. curtum
P. elegantissimum
P. metananum
P. mediocre
P. melo
P. penitum
P. thulesense

Penta p. bipatens P. archiovatum P. affine

P. bipes

P. granii
P. glyptopterum
P. incognitum
P. unipes

Hexa p. roseaceum P. cruciferum

P. incertum

Protoperidinium antarcticum (A.Schimp. ex G.Karst.) Balech (1973: 21)

BASIONYM: Peridinium antarcticum A.Schimp. ex G.Karst. (Karsten, 1905: 131, pi. 19, figs 1-4)

SYNONYM: Peridinium depressum (Bailey) N.Peters (1928: 63, fig. 17d-f)

ADDITIONAL REFERENCES: Balech (1976: 46, fig. 32), Andreoli et al. (1995: 473, fig. 19)

Fig. 3.19c, d

description: Cells large, expanded, 80-140 pm long (excluding horns), 80-120 pm

diameter. Surface ornamentation spiculate. Apical horn to 30 urn long; antapical horns of
equal length, to 25 pm long. Cingulum slightly helical, comparatively narrow, non-incised,
bordered by narrow ribbed ridges. Narrow growth bands usually present between plates.
Tabulation lpa, 4', 3a, 6", 3c, 6s?, 5'", lp, 1""; 1' plate large and rhombic (orthoform); 2a
plate trapezoidal (quadra).

distribution: Southern Ocean, south of Australia; type locality, Antarctic Ocean,

"Challenger" Expedition (Karsten, 1905); Bransfield Strait (Priddle, 1985); Southern Ocean,
near South Georgia (Dodge & Priddle, 1987); Terra Nova Bay, Ross Sea (Andreoli et al.,
1995); Weddell Sea (Estrada & Delgado, 1990); offshore from Davis Station, East Antarctica

235
 DINOFLAGELLATES

(Archer et al, 1996a); southern Indian Ocean, 46-69°S, 75°E (Okamoto et al, 1986);
KERFIX, 50°40'S 68°25'E, Southern Ocean (Kopczyhska et al, 1998).

The largest of the Antarctic dinoflagellates, P. antarcticum is sometimes mistaken for

P.pseudoantarcticum. However, while the latter has a similar tabulation pattern (orthoform
with a quadra 2a), it is smaller, narrower, and has longer and more expanded antapical horns.

Protoperidinium applanatum (Mangin) Balech (1974: 59)

BASIONYM: Peridinium applanatum Mangin (1926: 79, fig. 58)

SYNONYM: Peridinium obovatum E.J. F.Wood (1954: 242, fig. 159)

additional reference: Balech (1976: 69, fig. 50)

Fig. 3.19b

description: Cells transversely ellipsoidal, 23-45 urn long (excluding apical and antapical
horns), 43-60 um long overall, 30-55 urn diameter. Apical horn conical, 9-30 um long;
antapical horns equal in length, 1 1-25 um long. Cingulum c. 5 um wide, slightly displaced,
non-incised; cingular lists reduced; posterior list increasing in width to form the sulcal list.
Tabulation lpa, 4', 3a, 7", 6c, 5'", lp, 1"". Plate configuration: metaform quadra.

distribution: Southern Ocean, 44-63°S, south of Australia; type locality, Weddell Sea,
"Scotia" Expedition (Mangin, 1926); Bransfield Strait (Priddle, 1985); South Georgia
(Dodge & Priddle, 1987); Terra Nova Bay (Andreoli et al, 1995); Weddell Sea (Estrada &
Delgado, 1990); Southern Indian Ocean, 46°S 75°E (Okamoto et al, 1986).

Protoperidinium archiovatum (Balech) Balech (1974: 54)

BASIONYM: Peridinium archiovatum Balech (1958a: 84, pi. 3, figs 45-48)

additional REFERENCE: Balech (1976: 44, fig. 30)

description: [after Balech (1976)] Cells lenticular, lacking antapical horns. Tabulation lpa,
4', 2a, 6", 6c, 5'", lp, 1""; 6" plate very large and partially overlapping 5"; 2 apical
intercalary plates are present with 2a being very large. Plate configuration: metaform penta.

distribution: Type locality, Antarctic Peninsula (Balech, 1958a); Ross Sea (Watanabe,
1982); Terra Nova Bay, Ross Sea (Andreoli et al, 1995).

A member of the subg. Archaeoperidium (Balech, 1976), this uncommon species was not
seen by us. It resembles Preperidinium meunierii (Kolkwitziellaceae), but it has a clear
protoplasm, and it lacks a crescent-shaped sulcus and large left sulcal lists (Balech, 1976, as
Diplopeltopsis minor).

Protoperidinium areolatum (N.Peters) Balech (1974: 66)

BASIONYM: Peridinium areolatum N.Peters (1928: 33, fig. 8a-e)

ADDITIONAL REFERENCES: Bunt & Wood (1968), Balech (1976: 83, fig. 62)

description: [after Balech (1976)] Cells large, with well-developed apical and antapical
horns. Cingulum almost circular and slightly ascending. Strong surface reticulation on
thecae. Tabulation lpa, 4', 3a, 7", 6c, 5'", lp, 1"". Plate configuration: paraform quadra.

distribution: Type locality, Weddell Sea (Peters, 1928); Ross Sea (Watanabe, 1982).

Rarely reported and not seen by us.

Protoperidinium bellulum (Balech) Balech (1974: 59)

BASIONYM: Peridinium bellulum Balech (1971: 127, pi. 23, figs 407-41 1)

additional reference: Balech (1976: 67, fig. 49)

Fig. 3.20a, b

236
 DINOFLAGELLATES

Figure 3.20. Protoperidinium species, a, b, P. bellulum, SEM Oblique ventral and dorsal
views,c, d, P. charcoti, SEM Ventral and dorsal views. Scale bars: a-d = 10 um.

237
 DINOFLAGELLATES

Figure 3.21. Protoperidinium species, a, b, P. defectum, SEM Ventral and dorsal views, c,

d, P.elegantissimum, SEM Oblique dorsal and apical views. Scale bars: a-d = 10 urn.

238
 DINOFLAGELLATES

description: Cells transversely ellipsoidal, 25-36 um long (excluding apical horn and
spines), 34-42 urn overall, 28-36 urn diameter. Apical horn conical, 3 um long, expanded at
the tip. Hypotheca with 2 antapical horns set rather far apart, c. 7 um long; left antapical
horn slightly shorter that the right. Cingulum 5 um wide, non-incised, displaced by 1

cingular width; cingular lists greatly reduced, ribbed. Sulcus extending slightly into epitheca,
widening posteriorly towards antapex. Tabulation lpa, 4', 3a, 7", 6c, 5'", lp, 1"". Plate
configuration: metaform quadra.

distribution: Southern Ocean, 62°S, south of Australia; type locality, Drake Passage
(Balech, 1971); Ross Sea (Watanabe, 1982); Bransfield Strait (Priddle, 1985); Southern
Ocean, near South Georgia (Dodge & Priddle, 1987); Terra Nova Bay, Ross Sea (Andreoli et
al, 1995).

Protoperidinium bipatens Balech (1973: 17, pi. 4, figs 62-76)

ADDITIONAL REFERENCES: Balech(1974: 54; 1976: 44, fig. 31)

description: [after Balech (1973)] Antapical horns well developed, with acute spines at
their tips. Sulcus wide at the top (near the cingulum), narrowing close to the antapex and
expanding below this. Tabulation lpa, 4', 2a, 6", 6c, 5'", lp, 1""; 6" plate very large and
partially overlapping 5"; 2 apical intercalary plates present, with 2a being very large. Plate
configuration: orthoform penta.

distribution: Type locality, Bellingshausen Sea (Balech, 1973); Southern Ocean, near
South Georgia (Dodge & Priddle, 1987); Ross Sea (Watanabe, 1982).

A member of subg. Archaeoperidium (Balech, 1976), this uncommon species was not seen
by us.

Protoperidinium charcoti (Balech) Balech (1974: 60)

BASIONYM: Peridinium charcoti Balech (1958b: 394, pi. 3, figs 79-90)

ADDITIONAL REFERENCE: Balech (1976: 60, fig. 43)

Fig. 3.20c, d

DESCRIPTION: Cells almost pentagonal, 60-70 um long, 80-90 um wide. Sides of the epitheca
strongly concave; sides of the hypotheca convex. Apical horn prominent, narrow, the 2
antapical horns well developed and pointed. Cells with a noticable central bulge. Tabulation
lpa, 4', 3a, 7", 6c, 5'", lp, 1"". Plate configuration: metaform quadra.

distribution: Prydz Bay, East Antarctica; type locality, Terre Adelie (Balech, 1958b); Terra
Nova Bay, Ross Sea (Andreoli et al., 1995).

Protoperidinium concavum (Mangin) Balech (1974: 60)

BASIONYM: Peridinium concavum Mangin (1926: 79, figs 21, II)

SYNONYMS: Peridinium granii auct. non Ostenfeld (1906), sensu Peters (1928: fig. 13a-f)
Peridinium mite auct. non Pavillard (1916), sensu Wood (1954: 238, fig. 1 17b, c, non fig. 117a)
Peridinium petersii Balech (1958a: 88, pi. 5, figs 100-108)

ADDITIONAL REFERENCE: Balech (1976: 63, fig. 44)

DESCRIPTION: [after Balech (1976)] Cells large, 57-119 um long, 57-127 um diameter. Sides
of the epitheca strongly concave; sides of the hypotheca convex. Apical horn narrow;
antapical horns reduced to 2 short spines. Tabulation lpa, 4', 3a, 7", 6c, 5'", lp, 1"". Plate
configuration: metaform quadra.

distribution: Terra Nova Bay, Ross Sea (Andreoli et al., 1995); Ross Sea (Watanabe,
1982); Antarctic Peninsula (Mangin, 1926).

239
 DINOFLAGELLATES

Protoperidinium cf. crassipes (Kofoid) Balech (1988: 1 10, pi. 43, figs 5-7)
SYNONYM: Peridinium crassipes auct. non Kofoid, sensu Peters (1928: 42, fig. 1
1)

additional REFERENCE: Balech (1976: 60, fig. 42)

description: [after Balech (1976)] Cells stout and rather large, 77-97 urn long, 83-99 urn
diameter. Cingulum not deeply excavated; lists with broken ribs. Thecae ornamented with a
strong reticulum and abundant irregularly distributed pores. Tabulation lpa, 4', 3a, 7", 6c,
5'", lp, 1"". Plate configuration: metaform quadra. The 3'" plate is very asymmetrical.

distribution: Ross Sea (Watanabe, 1982).

Uncommon and not seen by us. Balech (1976) noted that this species is very close to some of
the P.crassipes auct., but it is clearly different from Peridinium crassipes Kofoid,
subsequently transferred to Protoperidinium by Balech (1988). A new name is not proposed
until a study of the crassipes group has been completed.

Protoperidinium cruciferum (Balech) Balech (1973: 25, pi. 7, figs 134-137)

BASIONYM: Peridinium cruciferum Balech (1971: 131, pi. 23, figs 421^28)

ADDITIONAL REFERENCE: Balech (1976: 79, fig. 59)

description: Cells subspherical, 26.5-32.5 urn long (excluding horns), 26.5-32.5 urn
diameter. Thecae with a surface pattern of small crosses; epitheca with a small apical horn;
hypotheca with short (c. 3 urn) antapical spines. Cingular lists ribbed. Tabulation lpa, 4', 3a,
7", 6c, 5'", lp, 1"". Plate configuration: metaform hexa.
distribution: Bellingshausen Sea (Balech, 1973); Bransfield Strait (Priddle, 1985);
Southern Ocean, near South Georgia (Dodge & Priddle, 1987); KERFIX, 50°40'S 68°25'E,
Southern Ocean (Kopczyhska et al., 1998); Ross Sea (Watanabe, 1982).

Protoperidinium curtum (Balech) Balech (1973: 22, pi. 6, figs 103-1 1 1)

BASIONYM: Peridinium curtum Balech (1958b: 393, pi. 3, figs 62-70)

SYNONYM: Peridinium ovatum auct. non Pouchet (1883), sensu Wood (1954: 236, fig. 1 12c, d)

additional REFERENCE: Balech (1976: 69, fig. 51)

Fig. 3.22d

description: Cells transversely ellipsoidal, 35-62 urn long, 30-60 urn diameter. Thecae
smooth, with scattered pores; epitheca with a short conical apical horn; hypotheca with
2 short divergent antapical horns. Cingulum c. 4 urn wide, non-incised, displaced by 1

cingular width; cingular lists broad, projected laterally, irregularly ribbed. Sulcus extending
posteriorly to antapex, but only slightly towards apex. Tabulation; lpa, 4', 3a, 7", 6c, 5'", lp,
1"". Plate configuration: metaform quadra.

distribution: Bellingshausen Sea (Balech, 1973); Southern Ocean, 50-63°S, south of

Australia; Southern Ocean, near South Georgia (Dodge & Priddle, 1987); Ross Sea
(Watanabe, 1982); Antarctic and sub-Antarctic waters (Hara & Tanoue, 1985).

Protoperidinium defectum (Balech) Balech (1974: 53)

BASIONYM: Peridinium defectum Balech, in Balech & El-Sayed (1965: 1 18, pi. 3, figs 45-50)

ADDITIONAL REFERENCES: Balech (1976: 42, fig. 29), Andreoli et al. (1995: 473)

Fig. 3.21a, b

DESCRIPTION: Cells subspherical, 14-29 urn long (excluding horns), 15-25 urn diameter. Cell
surface ornamented with large pores that are mostly concentrated around the cingulum.
Epitheca with a well-developed apical horn 3.6-10 urn long. Hypotheca with 2 well-
developed narrow antapical horns 1.3-10 urn long; left horn always longer than the right.

240
 DINOFLAGELLATES

Figure 3.22. Protopehdinium species, a, P. glyptopterum, SEM Apical view, b, P. granii,

SEM Oblique apical view, c, P. incertum, TEM Apical view, d, P. curtum, TEM Apical
view. Scale bars: a-d = 10 |im.

241
 DINOFLAGELLATES

Figure 3.23. Protoperidinium species, a. b, P. incognitum. a. SEM Ventral view. b. SEM

Apical view, c, d, P. latistriatum. c, SEM Oblique dorsal view, d, SEM Antapical view.
Scale bars: a-d = 10 um.

242
 DINOFLAGELLATES

Cingulum slightly displaced at the sulcus, broad, to 25% of cell length, non-incised; cingular
listsnarrow, regularly ribbed. Tabulation lpa, 4', 2a, 6", 3c, 6s?, 5'", lp, 1""; 6" plate
extremely large and partly overlapping 5"; in ventral view the 6" plate covers most of the
epitheca and distorts the sulcus. Plate configuration: paraform quadra.

distribution: Type locality, Weddell Sea (Balech & El-Sayed, 1965); Widespread in the
Nova Bay (Andreoli et al., 1995).
Antarctic; Terra

A member of subg. Archaeoperidium (Balech, 1976), the small size, long apical and
antapical horns, and the very large 6" plate are very distinctive. Protoperidinium defectum
resembles P. minusculum Pavill. from Northern Hemisphere waters.

Protoperidinium elegantissimum (Balech) Balech (1974: 60)

BASIONYM: Peridinium elegantissimum Balech (1958a: 89, pi. 5, figs 109-1 15)

additional reference: Balech (1976: 67, fig. 48)

Fig. 3.21c, d

description: Cells transversely ellipsoidal, 53-59 \im long (excluding horns), 40-56 urn
diameter. Thecae smooth, with scattered pores; epitheca slightly concave, tapering to the
apical horn; hypotheca with 2 almost parallel antapical horns. Cingulum slightly incised;
cingular lists narrow, irregularly ribbed. Tabulation; lpa, 4', 3a, 7", 6c, 5'", lp, 1"". Plate
configuration: metaform quadra.

distribution: Southern Ocean, 55-62°S, south of Australia; type locality, Terre Adelie
(Balech, 1958a); Bransfield Strait (Priddle, 1985); Ross Sea (Watanabe, 1982).

Protoperidinium glyptopterum Balech (1973: 25, pi. 7, figs 123-133)

ADDITIONAL REFERENCES: Balech (1974: 62; 1976: 75, fig. 55)

Fig. 3.22a

description: Cells transversely ellipsoidal, 56-65 |im long, 65-72 urn diameter. Thecal
surfaces smooth, with scattered pores; epitheca with a short narrow apical horn; hypotheca
with 2 short antapical spines. Cingulum 5 \im wide, non-incised, displaced by 1 cingular
width; cingular lists reduced, ribbed. Sulcus extended anteriorly barely into epitheca but
posteriorly into hypotheca to antapex; sulcal lists well developed. Tabulation; lpa, 4', 3a, 7",
6c, 5"', lp, 1"". Plate configuration: metaform penta.
distribution: Type locality, Bellingshausen Sea (Balech, 1973); Southern Ocean, 50-63°S,
south of Australia; Southern Ocean, near South Georgia (Dodge & Priddle, 1987); Ross Sea
(Watanabe, 1982).

Protoperidinium granii (Ostenf.) Balech (1974: 65)

BASIONYM: Peridinium granii Ostenf. (Ostenfeld, 1906: 15)

ADDITIONAL REFERENCES: Dodge (1982: 196, fig. 23A), Balech (1988: 107, pi. 40, figs 9-12)

Fig. 3.22b

description: Cells subrectangular, 49-80 urn long, 49-56 \xm diameter. Thecal surfaces
with fine reticulations or scattered pores; epitheca with straight to convex sides and a short
stout apical horn; hypotheca with straight to concave sides, with 2 short divergent antapical
spines. Cingulum 5 u.m wide, slightly incised and displaced; cingular lists regularly ribbed.
Sulcus extending anteriorly barely into epitheca but posteriorly into hypotheca to antapex;
sulcal lists well developed. Tabulation lpa, 4', 3a, 7", 6c, 5'", lp, 1"". Plate configuration:
metaform penta.
distribution: Southern Ocean, 56-60°S, south of Australia; type locality, Faeroe Is.

(Ostenfeld, 1906); South Atlantic at 47°S (Balech, 1988).

243
 DINOFLAGELLATES

Protoperidinium incertum (Balech) Balech (1973: 26, pi. 7, figs 138-141)

BASIONYM: Peridinium incertum Balech (1958a: 91, pi. 6, figs 133-140)

ADDITIONAL REFERENCES: Balech (1974: 67; 1976: 81, fig. 60)

Fig. 3.22c

description: Cells pyriform to almost pentagonal, 44-62 um long (excluding horns), 50-71 urn
long overall, 47-59 urn diameter. Thecal surfaces smooth; epitheca straight, concave towards
a short stout apical horn; hypotheca with divergent antapical spines to c. 10 um long.
Cingulum non-incised, displaced by 0.5-1 cingular width; cingular lists ribbed. Sulcus not
extending to antapex or into epitheca; sulcal lists forming 2 false spines at antapex (Balech,
1976). Tabulation lpa, 4', 3a, 7", 6c, 5'", lp, 1"" (2a hexa, large). Plate configuration:
metaform hexa.
distribution: Type locality, South Sandwich Is. (Balech, 1958a); Southern Ocean, 44°S,
south of Australia; Terra Nova Bay (Andreoli et al., 1995); Ross Sea (Watanabe, 1982).

Protoperidinium incognitum (Balech) Balech (1974: 62)

BASIONYM: Peridinium incognitum Balech (1959: 24, pi. 2, figs 43-45)

ADDITIONAL REFERENCE: Balech (1976: 77, fig. 56)

Fig. 3.23a, b

description: Cells subspherical, c. 40 um long (excluding horns), c. 38 um diameter. Thecal

surfaces strongly reticulated and with scattered pores; epitheca with convex sides and a short
stout apical horn supported by a septa along the apical plate boundaries; hypotheca convex,
with 2 divergent antapical spines 6 um long, with membranes. Cingulum 5 um wide, slightly
incised and displaced, but without an overhang; cingular lists broad, regularly ribbed. Sulcus
extending anteriorly barely into epitheca but posteriorly into hypotheca to antapex; sulcal
lists well developed. Tabulation lpa, 4', 3a, 7", 6c, 5'", lp, 1"" (2a plate is very large). Plate

configuration: metaform penta.

distribution: Southern Ocean, 56-60°S, south of Australia; type locality, Antarctic

Peninsula (Balech, 1959); Antarctic and sub-Antarctic waters (Balech, 1976); Southern
Ocean, near South Georgia (Dodge & Priddle, 1987); Ross Sea (Watanabe, 1982).

Protoperidinium latistriatum (Balech) Balech (1974: 57)

BASIONYM: Peridinium latistriatum Balech (1958a: 86, pi. 4, figs 66-80)

SYNONYM: Peridinium subinerme auct. non Paulsen (1904), sensu Peters (1928: 52, fig. 15) pro parte

ADDITIONAL REFERENCE: Balech (1976: 50, fig. 35)

Fig. 3.23c, d

description: Cells pentagonal, 30-112 um long, 30-97 um diameter. Thecal surfaces with
pores, reticulations and minute spines. Epitheca with slightly convex sides; apical horn
greatly reduced, with an apical pore. Hypotheca rounded, with greatly reduced horns.
Cingulum 4 um wide, incised, equatorial, slightly displaced; surface with faint reticulations
and pores; cingular lists narrow, ribbed. Sulcus barely extending into epitheca, but
broadening posteriorly into hypotheca; sulcal lists very narrow. Tabulation lpa, 4', 3a, 7", 6c,
5'", lp, 1"". Plate configuration: orthoform quadra. Growth bands to 5 um wide, usually

present between plates.

Distribution: Southern Ocean, 51-62°S, south of Australia; type locality, Antarctic

Peninsula (Balech, 1958a); Southern Ocean, near South Georgia (Dodge & Priddle, 1987).

Specimens found in the Southern Ocean, south of Australia were considerably smaller than
those reported by Balech, and they had minute spines on the thecal surfaces.

244
 DINOFLAGELLATES

Figure 3.24. Protoperidinium species, a, b, P. mediocre, a, SEM Oblique ventral and apical
views, b, SEM Dorsal view, c, P. melo, SEM Dorsal view, d, P. penitum, SEM Dorsal view.
Scale bars: a-d = 10 urn.

245
 DINOFLAGELLATES

Figure 3.25. Protoperidinium metananum. a, SEM Ventral view; some plates dislodged, b,
SEM Ventral view, c, SEM Oblique apical view, d, SEM Oblique antapical view. Scale bars:
a-d = 10 urn.

246
 DINOFLAGELLATES

Protoperidinium mediocre (Balech) Balech (1974: 61)

BASIONYM: Peridinium mediocre Balech (1958b: 389, pi. 2, figs 54-61)

ADDITIONAL REFERENCES: Balech (1976: 65, fig. 46), Andreoli et al. (1995: 473)

Fig. 3.24a, b

description: Cells subspherical, 30-40 um long (excluding horns), 30-40 um diameter.

Thecal surfaces smooth, with scattered pores; epitheca tapering to a short apical horn
supported by septa along apical plate boundaries; hypotheca with 2 small close-set antapical
horns that incline inwards. Cingulum c. 4 um wide, non-incised, ornamented with pores,
displaced by cingular width; cingular lists wide, irregularly ribbed. Tabulation lpa, 4', 3a, 7",
1

3c, 6s?, 5'", lp, 1"". Plate configuration: pentagonal (metaform); 2a plate rhombic (quadra).

distribution: Widespread in oceanic and coastal areas of the Antarctic; type locality, Terre
Adelie (Balech, 1958b); Southern Ocean, near South Georgia (Dodge & Priddle, 1987); Ross
Sea (Andreoli et al, 1995).

This species can be distinguished from most others with a similar plate pattern by its almost
spherical shape and small, close-set horns. It is most similar to P. concavum, but the horns of
that species extend outwards, whereas those of P. mediocre slope inwards.

Protoperidinium melo (Balech) Balech (1974: 61)

BASIONYM: Peridinium melo Balech (1971: 121, pi. 22, figs 392-399)

ADDITIONAL REFERENCES: Balech (1976: 73, fig. 53), Andreoli et al. (1995: 473)

Fig. 3.24c

DESCRIPTION: Cells subspherical, 30-42 um long (excluding horn and spines), 35-45 um
diameter. Thecal surfaces smooth, with scattered pores and irregular faint spines; epitheca
tapering to an apical horn; hypotheca with 2 small parallel antapical spines. Cingulum c. 5 um
wide, non-incised, ornamented with pores; cingular lists broad, irregularly ribbed. Tabulation
lpa, 4', 3a, 7", 6c, 5'", lp, 1"". Plate configuration: metaform quadra.

distribution: Southern Ocean, 53-63°S, south of Australia; type locality, Drake Passage
(Balech. 1971); Southern Ocean, near South Georgia (Dodge & Priddle, 1987); Terra Nova
Bay (Andreoli et al, 1995); Ross Sea (Watanabe, 1982).

Protoperidinium metananum (Balech) Balech (1974: 61)

BASIONYM: Peridinium metananum Balech, in Balech & El-Sayed (1965: 120, pi. 2, figs 27-33)

ADDITIONAL REFERENCE: Balech (1976: 58, fig. 41)

Fig. 3.25a-d

description: Cells spherical, 23-34 um long (excluding horns), 25-36 um diameter. Thecal
surfaces reticulate, with scattered pores; epitheca with convex sides, tapering to a short
apical horn; hypotheca with 2 short close-set antapical horns. Cingulum c. 5 um wide, non-
incised; surface reticulate, with pores; cingular lists narrow, irregularly ribbed. Sulcus
extending slightly into epitheca; sulcal lists wide. Tabulation lpa, 4', 3a, 7", 6c, 5'", lp, 1"".
Plate configuration: metaform quadra.
distribution: Southern Ocean, 44-64°S, south of Australia; Antarctic waters (Balech, 1976,
1988; Hara & Tanoue, 1985).

Protoperidinium penitum (Balech) Balech (1974: 61)

BASIONYM: Peridinium penitum Balech (1971: 1 19, pi. 22, figs 400-406)
SYNONYM: Peridinium ovatum auct. non Pouchet (1883), sensu Peters (1928: 40, fig. 10)

ADDITIONAL REFERENCE: Balech (1976: 72, fig. 52)

247
 DINOFLAGELLATES

Fig. 3.24d

description: Cells subspherical, 30-42 nm long (excluding horn and spines), 37-48 um
long overall, 29-50 um diameter. Thecal surfaces smooth, with scattered pores and a row of
rimmed pores along cingular margin of epitheca and hypotheca; epitheca with a long broad
apical horn; hypotheca with 2 small antapical spines, the left longer than the right. Cingulum
c. 6 um wide, non-incised, displaced by 1 cingular width; surface with scattered pores;

cingular lists narrow, irregularly ribbed. Sulcus barely extending into epitheca, extending
posteriorly to antapex; sulcal lists wide, left list highly developed to form a false spine.
Tabulation lpa, 4', 3a, 7", 6c, 5'", lp, 1"". Plate configuration: metaform quadra.

DISTRIBUTION: Southern Ocean, 60-63°S, south of Australia; type locality, Drake Passage
(Balech, 1971); Terra Nova Bay (Andreoli et al, 1995); Southern Ocean, near South Georgia
(Dodge & Priddle, 1987); Ross Sea (Watanabe, 1982).

Protoperidinium rosaceum (Balech) Balech (1974: 59)

BASIONYM: Peridinium rosaceum Balech (1958a: 87, pi. 4, figs 81-85, 88-91)

additional reference: Balech (1976: 50, fig. 36)

Fig. 3.26a

description: Cells kite-shaped, with the epitheca and hypotheca having straight sides, 50-80
um long, 48-69 um diameter. Thecal surfaces reticulate, with scattered pores; epithecal sides
meet at the apical pore with no apparent apical horn; hypotheca with an almost flat base and
no antapical horns or spines. Cingulum relatively wide (5 um), excavated and faintly
reticulated; cingular lists narrow. Sulcus extending only slightly into the epitheca, round in
shape, increasing in width posteriorly; sulcal lists ribbed, narrow. Sutures comparatively
wide (2 um), with striae. Tabulation lpa, 4', 3a, 7", 6c, 5'", lp, 1"". Plate configuration:
orthoform hexa.

distribution: Type locality, Terre Adelie (Balech, 1958a); Southern Ocean, near South
Georgia (Dodge & Priddle, 1987); Ross Sea (Watanabe, 1982).

Protoperidinium thulesense (Balech) Balech (1973: 27)

BASIONYM: Peridinium thulesense Balech (1958a: 92, pi. 6, figs 152-160)

SYNONYMS: Peridinium conicum f. islandica Braarud (1935: 108, fig. 27)

Peridinium sympholis Hermos. & Balech (Hermosilla & Balech, 1969: 9, figs 1-3)

additional references: Balech (1976: 56, fig. 39), Dodge (1982: 202, fig. 23L, N), Balech
(1988: 105, pi. 39, figs 1-5)

Fig. 3.26c, d

description: Cells pentagonal to polygonal, 32-40 um long, 34-40 um diameter. Thecal

surfaces reticulate, with scattered pores; epitheca with slightly concave sides and a small
apical horn that is supported by septa along the thecal plate boundaries; hypotheca with an
almost flattened base and no apparent antapical spines. Cingulum incised, displaced 0.5-1
cingular width; cingular lists absent. Sulcus narrow and straight, extending posteriorly to
antapex, but not into the epitheca; sulcal lists not apparent. Tabulation lpa, 4', 3a, 7", 6c, 5'",
lp, 1"". Plate configuration: metaform quadra.
distribution: Type locality, South Sandwich Is. (Balech, 1958a); Southern Ocean, south of
Australia; Southern Ocean, near South Georgia (Dodge & Priddle, 1987); bipolar (Balech,
1976).

Protoperidinium unipes (Balech) Balech (1974: 63)

BASIONYM: Peridinium unipes Balech (1962: 31, pi. 2, figs 46-56)

ADDITIONAL REFERENCE: Balech (1976: 77, fig. 57)

Fig. 3.26b

248
 DINOFLAGELLATES

Figure 3.26. Protoperidinium species, a, P. rosaceum, SEM Ventral view, b, P. unipes, SEM
Apical view, c, d, P. thulesense. c, SEM Ventral view, d, SEM Apical view. Scale bars: a-d
= 10 urn.

249
 DINOFLAGELLATES

description: Cells pyriform, 33-65 urn long (excluding horn and spines), 28-120 um
diameter. Thecal surfaces reticulate, with scattered pores. Epitheca straight to convex,
tapering to a long broad apical horn. Hypotheca convex, the 2 antapical spines 12-14 um
long, equal, or the left spine slightly longer; membranes present on spines. Cingulum 5 um
wide, non-incised, displaced by 0.5 cingular width; surface reticulated; cingular lists broad,
ribbed. Sulcus extending slightly into epitheca and posteriorly to antapex; sulcal lists well
developed. Tabulation lpa, 4', 3a, 7", 6c, 5'", lp, 1"". Plate configuration: metaform penta.

distribution: Southern Ocean, 44-62°S, south of Australia; type locality, Antarctic

Peninsula (Balech, 1962).

Cells found in the Southern Ocean, south of Australia were smaller (33^40 um long) but
otherwise identical to those reported by Balech (1976).

250
 4. SILICOFLAGELLATES
Gustaaf M. Hallegraeff

10-70 urn in diameter. Several stages of the life

Silicoflagellates are yellow, unicellular protists
history of Dictyocha speculum Ehrenb. have been described following ultrastructural
examination (Moestrup & Thomsen, 1990) and growth studies of cultured material (Henriksen
et ai, 1993). These stages include skeleton-bearing unicells (uniflagellate), large spherical
cells, uninucleate naked cells (uniflagellate), an amoeboid (transition) stage and large
multinucleate cells (with one or more flagella).

The most common form is the skeleton-bearing cell with its characteristic basket-shaped,
siliceous skeleton. The skeleton consists of a polygonal basal ring with a central arch and
protruding radial, tubular silica elements or spines. Skeleton-bearing cells are essentially
biflagellate, although one flagellum is not always obvious, being represented only by its
basal body. The cytoplasm is very fragile and contains 30-50 yellow chloroplasts around the
periphery. Traditionally their taxonomy has been based exclusively on the morphology of the
siliceous skeleton (Gemeinhardt, 1930), but the validity of this approach has been questioned
due to the observed plasticity in skeleton configuration in clonal cultures (van Valkenberg,
1980; Henriksen et al., 1993). Silicoflagellates without the skeleton, the naked stage, have
also been described (van Valkenberg &
Norris, 1970; Henriksen et ai, 1993). Cells of the
naked form can be distinguished from other golden brown flagellates (such as the
raphidophyte Heterosigma) by the presence of a single anterior flagellum and the absence of
a trailing flagellum.

Concepts of species and genus were discussed by Sournia (1986), Moestrup & Thomsen
(1990) and Henriksen et al. (1993). As noted by Moestrup & Thomsen (1990), the widely
used generic name Distephanus Stohr (1880) is preoccupied by Distephanus Cassini (1817),
a member of the flowering plant family Asteraceae, but it sometimes still appears in the
phytoplankton literature.

Silicoflagellates occur in all oceans. Generally regarded as having low overall abundance in
the present-day seas, the skeleton-bearing stages are plentiful in the fossil record
(e.g. Deflandre, 1950; Loeblich et al, 1968) and have been used as a tool for the study of
Tertiary Antarctic climates (Mandra, 1969). Three extant species are recognised: Dictyocha
speculum is usually dominant in Atlantic and sub-Antarctic waters, with D. fibula more

common around the Subtropical Convergence (Van der Spoel et al., 1973; Van der Spoel,
1987). Dictyocha octonaria appears to be rare in Antarctic waters.

Order Dictyochales

Family Dictyochaceae

Dictyocha fibula Ehrenb. (Ehrenberg, 1837: 61, pi. 18, figs 54, 55; pi. 20, fig. 45; pi. 21,
fig. 42b; pi. 22, fig. 42c)

ADDITIONAL REFERENCES: van Valkenberg & Norris (1970), van Valkenberg (1980: 335, figs 1-20)

Fig. 4.1a, b

description: Cells uniflagellate. Silica skeleton cap-shaped, comprising a large rectangular

or rhombic ring, with 4 large spines radiating from the corners of the ring. Connecting bars
emanate from centre of each side of ring forming 4 windows. Smaller spines sometimes
projecting outwards from the large ring.

Class Dictyochophyceae or Order Silicoflagellida (of Subphylum Heterokonta).

251
 SILICOFLAGELLATES

Figure 4.1. Dictyocha species, a, b, D. fibula, a, SEM Solitary cell, b, SEM Siliceous skeletal
ring with projecting spines, c, d, D. octonaria. c, SEM Siliceous skeletal ring with projecting
spines, d, SEM Siliceous skeletal ring with projecting spines. Scale bars: a-d = 10 urn.

252
 SILICOFLAGELLATES

Figure Dictyocha speculum, a, LM Solitary cell, b, SEM Solitary cell, c, SEM Full
4.2.
silica of vegetative phase and attached cytoplasm, d, SEM Skeleton of cell
skeleton
undergoing division, e, SEM Internal view of interlocking skeletal ring, f, SEM External
view of an interlocking skeletal ring. Scale bars: a-f = 10 urn.

253
 SILICOFLAGELLATES

distribution: Southern Ocean between Australia and Antarctica.

Another 4-spined cell, Dictyocha antarctica Lohmann, has been reported from the South
Atlantic (Throndsen, 1996), but its radiating spines are significantly smaller.

Dictyocha octonaria Ehrenb. (Ehrenberg, 1844: 57)

SYNONYMS: Distephanus speculum var. octonarius (Ehrenb.) Jorg. (Jorgensen, 1899)
Octactis pulchra J.Schiller (1925b: 66)

ADDITIONAL REFERENCE: Throndsen (1996: 632, pi. 6)

Fig. 4.1c, d

description: Cells uniflagellate. Silica skeleton to 70 um diameter (distance between the

tips of the longest spines), characterised by an outer silica ring c. 25 urn diameter, with 8
radiating skeletal elements protruding as spines, 2 spines usually slightly longer than the
others, and a very delicate inner ring c. 18-24 urn diameter.

distribution: Coastal waters near the Vestfold Hills, East Antarctica.

Dictyocha speculum Ehrenb. (Ehrenberg, 1837: 61, pi. 18, figs 5-7)

SYNONYMS: Distephanus speculum (Ehrenb.) Haeckel (1887)

Cannopilus calyptra Haeckel (1887)

ADDITIONAL REFERENCES: van Valkenberg (1980: 335), Moestrup & Thomsen (1990: pi. 1-16),
Henriksen et al. (1993: 29, figs 3-11)

Fig. 4.2a-f

description: Cells uniflagellate. Silica skeleton to 70 um diameter (distance between the

tips of the longest spines), comprising 2 hexagonal rings of different sizes connected by 6
bars and with 6 conspicuous protruding spines; cytoplasm contained between the hexagonal
rings. Bars attached to corners of a small ±circular ring. Large hexagonal ring c. 17-23 um
diameter; small hexagonal ring c. 7-1 1 um diameter. Spines radiating from the corners of the
large ring c. 15 um long; smaller spines (usually 1-3) projecting from the small ring.
Flagellum located near one radial spine, c. 25 um long, covered in flagellar hairs. Protoplast
with a large central nucleus surrounded by numerous discoid golden brown chloroplasts and
other organelles.

distribution: Drygalski Glacier, Ross Sea and coastal waters off Davis Station, East
Antarctica; common in the Southern Ocean (Hada, 1970); Weddell Sea (Balech & El-Sayed,
1965; Fryxell, 1989); Weddell-Scotia Confluence (Garrison et al., 1987).

This silicoflagellate is one of the most spectacular and readily identified organisms in the
Southern Ocean. A small percentage of skeletons can have morphological anomalies, with a
5-fold or a 7-fold rather than 6-fold symmetry (sometimes referred to as var. pentagonus
Lemmerm. and var. septanarius (Ehrenb.) Jorg., respectively). Twin cells with opposed and
partly interlocked skeletons represent dividing cells.

254
 5. HAPTOPHYTES: ORDER PRYMNESIALES*
Harvey J. Marchant, Fiona J. Scott and Andrew T. Davidson

Haptophytes are small, biflagellate and often solitary cells characterised by scales on the cell
body and the presence of a haptonema, a thread-like or bulbous appendage protruding from
near the base of the flagella. The morphology of the scales is the principal feature used in the
identification of species. Until recently, many species remained undiscovered owing to their
small size and delicate structure.

Cells typically contain 2 saucer-shaped, parietal chloroplasts; Golgi bodies and endoplasmic
reticulum can vary between species. Aspects of the taxonomy, life and
cycle, ultrastructure
ecology of the haptophytes were discussed in detail by Green & Leadbeater (1984). The
haptonema, its structure and possible functions were documented by Hibberd (1980) and
Inouye & Kawachi (1994).
Included here are a number of haptophytes, particularly of the genus Chrysochromulina,
which have been rarely reported from Antarctic waters, are of doubtful identity, or are
species of which only scales have been observed.

Order Prymnesiales

Cells are usually motile with 2 acronematic flagella and an obvious haptonema that is often
well developed and coiled. Species descriptions frequently include reference to the scale
types, the number of layers of scales covering the cell body, and the patterning on the
proximal (inner) and distal (outer) surfaces of the scales, usually visible by TEM. Several
species (e.g. of Chrysochromulina) are mixotrophic.

Family Phaeocystaceae

Phaeocystis antarctica G.Karst. (Karsten, 1905: 130, pi. 18, fig. 16)

ADDITIONAL REFERENCES: Kornmann (1955: 220, figs 1-6), Kayser (1970: 195), Richardson &
Whitaker (1979: 110), Chang (1984: 306, figs 1-7), Davidson & Marchant (1992b: 5, figs lb, c, 2,

3), Marchant & Thomsen ( 994: 211), Medlin et al. ( 1 994: 207)
1

Figs 5.1a-e, 5.2a-d

DESCRIPTION: Life Cycle: Phaeocystis has a complex life cycle including haploid microzoo-
spores, macrozoospores and non-motile cells. The two life cycle stages commonly observed
are biflagellate motile cells and palmelloid colonial cells. However, non-motile cells have
been described for other members of the genus (Kornmann, 1955; Kayser, 1970; Cariou
etal., 1994).

Motile cells: Cells solitary, biflagellate and with a haptonema, spherical to ovoid, 5-8 um
diameter. Flagella equal, 8-12 um long, bearing hair-points. Haptonema short, stiff, with a
bulbous tip, arising between the flagella but in a different plane from the flagellar bases. Cell
body with 2 layers of organic scales; scales of outer layer usually almost circular, 0.18 * 0.19
um, to 0.19 x 0.27 um diameter (Hallegraeff, 1983; Marchant & Davidson, unpublished
data), flat, with a perpendicular outwardly raised rim, usually with 45-48 ridges radiating
from a rectangular unornamented centre on both surfaces. Scales of inner layer with a
strongly inflexed rim, oval, 0.10 * 0.13 um diameter, with 30 ridges radiating from an oval
unornamented centre on both surfaces. Motile cells also release 'star' arrays. Threads of
these arrays are 20-100 um long (Buck & Garrison, 1983; Fryxell, 1989), composed of

Class Haptophyceae or Class Prymnesiophyceae.

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 HAPTOPHYTES: ORDER PRYMNESIALES

single or overlapping segments and tapering distally. Proximal ends of the threads overlap to
form a membrane-bound pentagon (Fig. 5.1e).

Colonial Colonies commonly spherical, from c. 10 urn diameter (containing only a few
cells:
cells) to 2 cm diameter (containing thousands of cells), comprising a mucilaginous matrix
with the cells arranged randomly around the periphery of the matrix. Cells spherical, 5-10 urn
diameter, lacking body scales, haptonema and flagella.

Other motile morphs: Ultrastructural studies indicate that there are at least two Phaeocystis-
like algae in Antarctic waters that are difficult to distinguish (Marchant & Davidson,
unpublished data). The three scale morphologies, all with distinct rims (Table 5.1; Fig. 5.2),
differ from the foregoing description off. antarctica.

Table 5.1. Scale morphologies of Phaeocystis-\ike morphs in Antarctic waters other than
those currently described for P. antarctica.

Morph Large scale Large scale patterning Small scale Small scale
diameter (fim) diameter (um) patterning

1 0.57 x 0.38 Radial on one side 0.24 x 0.19 Radial

2 0.22 x 0.19 Radial on one side; 0.16 x 0.13 unknown

central knob and
thickened annulus

3 0.34 x 0.26 Radial on one side 0.12 x 0.10 unknown

Phaeocystis pouchetti-Mke, lobed colonies with clumped cell distributions or cells aggregated at
one pole have also been observed in Antarctic waters (Marchant, unpublished data; Fig. 5. Id).

distribution: Circumpolar (Hart, 1942); Weddell Sea (Baumann et ai, 1994); Prydz Bay,
East Antarctica (Davidson & Marchant, 1987); coastal waters off Davis Station, East
Antarctica (Davidson & Marchant, 1992); Ross Sea (DiTullio et al., 2000)); type locality,
Antarctic Ocean, "Challenger" Expedition (Karsten, 1905); Bransfield Strait (Kang & Lee,
1995).

Phaeocystis antarctica was genetically differentiated as a distinct species by Medlin et al.

(1994) and is the only Phaeocystis species currently recognised from Antarctic waters.
However, no definitive characterisation of its morphology has been published. EM studies
(Davidson & Marchant, unpublished data) indicate a strong similarity with descriptions of
the flagellate stage of P. pouchetii (Har.) Lagerh. (Parke et ai, 1971), while the life cycle
and colony morphology resemble P. globosa Scherff. (Baumann et ai, 1994; Rousseau et al.,
1994). An additional, non-motile cell stage has been observed in cultured material
(Kornmann, 1955; Kayser, 1970).
Kornmann (1955) described the formation of non-motile cells from flagellate cells, which in
turn formed colonies. In contrast, Kayser (1970) reported that the non-motile cells became
attached to solid surfaces and released new, free, single cells and colonies into the water
column. These non-motile, single cells were proposed as being benthic in nature or attached
to particles. Whether or not both authors referred to the same cell type is uncertain.

Phaeocystis antarctica is among the most abundant of Antarctic phytoplankton species. Cell
7
concentrations can reach 6 x 10 cells per litre, giving the sea a brownish, oily appearance
and releasing sufficient quantities of dimethylsulphide (DMS) to produce a pungent smell.
It is one of the first algae to bloom in coastal Antarctica where it frequently dominates the

phytoplankton in the ice and in ice-edge waters during spring (Fryxell et ai, 1984; Perrin et
ai, 1987; Garrison & Buck, 1989).

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 HAPTOPHYTES: ORDER PRYMNESIALES

Figure 5.1. Phaeocystis antarctica. a, LM

Motile cells; note haptonema on the left-hand cell.
b, LMColonial cells, early stage, within a mucilaginous matrix, c, LM Colonial cells, d, LM
Large colony of non-motile cells, e, TEM 'Star' array threads of motile cells. Scale bars: a, b
= 5 urn; c = 20 um; d = 50 um; e = 1 um.

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 HAPTOPHYTES: ORDER PRYMNESIALES

Figure 5.2. Phaeocystis-like morphs. a, TEM motile cell showing features common to all
morphs, viz. flagella, haptonema (arrow) and body scales, b, TEM The 2 scale types of
motile cells (Morph 1). c, TEM Scale of motile cells (Morph 2). d, TEM Scale of motile cells
(Morph 3). Scale bars: a, b, d = 1 urn; c = 0.5 urn.

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 HAPTOPHYTES: ORDER PRYMNESIALES

Figure 5.3. Chrysochromulina acantha. a, TEM Shadow cast of collapsed cell, b, TEM Detail
of oval plate scales (pi) and spine scales (sp) of same cell, c, TEM Collapsed cell, d, TEM
Detail of oval plate scales and spine scales. Scale bars: a = 10 urn; b, c =
1 pm; d = 0.5 pm.

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 HAPTOPHYTES: ORDER PRYMNESIALES

Family Prymnesiaceae

Chrysochromulina
Cells have homodynamic flagella and a coiling haptonema, and electron microscopy is usually
required to identify the species. As well as being photosynthetic, these fragile haptophytes
ingest picoplanktonic and nanoplanktonic organisms by phagocytosis. Several species of
Chrysochromulina have been found in the Southern Ocean (Marchant, 1988), and a number of
Chysochromulina-Yike scales are included here to alert readers to possible new taxa.

Chrysochromulina acantha B.Leadb. & Manton (Leadbeater & Manton, 1971: 67,
figs 1-13)

additional REFERENCES: Hallegraeff (1983: 496, fig. 5a-c), LeRoi & Hallegraeff (2004: 75, fig. 2)

Fig. 5.3a-d

description: Cells saddle-shaped, ventrally concave, dorsally convex, 6-10 um long and
wide, biflagellate. Body scales of 2 types: outer layer of short spine scales, with spine c. 0.5
um long and attached to base-plate by 4 rigid arched supporting struts, the base-plate
elliptical to circular, concave, 0.45-0.5 pm diameter, with distal surface patterned with
delicate ±radiating striae; underlayer of flat oval plate scales c. 0.6 um long, with a slightly
thickened rim, patterned similar to spine scale base-plate. Flagella equal, c. 40 um long,
arising from concave ventral surface of cell c. one-third along cell length from posterior end.
Haptonema c. 40 um long when extended, coiled into 15-20 gyres when retracted.
distribution: Southern Ocean, south of Australia; Great Britain and Norway (Leadbeater &
Manton, 1971).

Figure 5.4. Chrysochromulina species, a, C. chiton, TEM (from Moestrup, 1979). b, C.

leadbeateri, TEM Plate scale. Scale bars: a, b = 200 nm.

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 HAPTOPHYTES: ORDER PRYMNESIALES

Chrysochromulina bergenensis B.Leadb. (Leadbeater, 1972a: 75, figs 1 (type), 47, 48)

ADDITIONAL REFERENCE: Manton & Leadbeater ( 1974: 9, figs 24-42)

DESCRIPTION: Cells ovoid, with a flattened flagellar pole and a slight central depression, 4 x 3.5
jam, biflagellate; body covered in unmineralised scales. Body scales of 2 types: outer layer of
large flat elliptical rimless scales, patterned with radiating striae on proximal face,
patternless on distal face; small plate scales forming an inner layer or positioned in
interstices of larger plates, elliptical, 0.4 * 0.32 urn, concave with an inflexed rim, patterned
as larger scales. Flagella 10-20 um long. Haptonema c. 8 urn long.

distribution: Southern Ocean, south of Australia; Norway and Mediterranean and Adriatic
Seas (Leadbeater, 1972b).

Chrysochromulina chiton Parke & Manton, in Parke et al. (1958: 225, figs 1-37)

ADDITIONAL REFERENCES: Leadbeater (1972b: pi. 4, fig. 28), Moestrup (1979: 68, fig. 10),
Hallegraeff (1983: 496, fig. 6), LeRoi & Hallegraeff (2004: 805, fig. 12)

Fig. 5.4a

DESCRIPTION: [after Parke et al. (1958)] Cells spherical to ovoid, with a flattened flagellar
pole depressed centrally across one axis, typically 5-8 um diameter, biflagellate; body
covered in unmineralised scales. Two types of body scales form a single layer over cell:
large scales elliptical, 2.4-2.9 um long, 1.9-2.42 um wide, saucer-shaped, with a broad rim
and radiating pattern of striae on proximal surface, distal surface patternless; small scales
circular to elliptical, 0.9-1.4 um long, 0.7-1.1 um wide, otherwise similar to large scales.
Flagella equal to subequal, to 25 um long, typically 2.5-3.5 times body length, arising from
slightly off-centre of depression. Haptonema c. 35 um long, typically 4-5 times body length
when extended.

DISTRIBUTION: Southern Ocean, south of Australia; New Zealand (Moestrup, 1979); northern
seas at 47°36'N 04°18'W (Parke et al, 1958).

Chrysochromulina ericina Parke & Manton, in Parke et al. (1956: 389, figs 1-19)

ADDITIONAL REFERENCES: Moestrup (1979: 68, figs 20-22), LeRoi & Hallegraeff (2004: 81, fig. 15)

Fig. 5.5a-c

description: Cells ovoid to oblong, slightly flattened, 5-12 um long, 4-10 um wide,
flagellar pole obliquely truncated, with a slight ±central depression, biflagellate; body
covered in spinous unmineralised scales. Body scales of 2 types: spine scales 28-30 long,
with a circular base-plate 1-1.4 um diameter, marked with concentric lines on distal surface,
with a spine 9-15 um long and abruptly truncate; plate scales numerous, elliptical, 0.6-0.9
um long, 0.5-0.7 um wide, with a pattern of fine striae on one surface and a slightly raised
rim. Flagella equal, c. 20 um long, 2-2.5 times body length, arising from slightly off-centre
of depression. Haptonema c. 53 um long, 4-5 times body length when extended.

distribution: Southern Ocean, south of Australia; New Zealand, and also widely distributed
in northern seas (Moestrup, 1979); 50°15'N 04°13'W (Parke et al., 1956).

Moestrup (1979) and Leadbeater (1972b) noted variability in the dimensions of the scales.

Chrysochromulina hirta Manton (1978: 13, figs 1-17; type figs 5, 7)

ADDITIONAL REFERENCES: Kawachi et al. (1991: 563, figs 1-29), LeRoi & Hallegraeff (2004: 83,
fig. 17)

Fig. 5.6a-d

description: Cells ovoid to spherical, c. 6 um diameter (dried preparation), biflagellate;

body covered in unmineralised scales. Body scales of 3 types: long spine scales distributed

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 HAPTOPHYTES: ORDER PRYMNESIALES

uniformly over cell, 20-30 um long, with spine basally forked and grooved and attached to
base-plate by 4 rigid supporting struts, the base-plate elliptical, concave, 2-2.5 um diameter,
the distal surface patterned with delicate ±radiating striae; short spine scales distributed
uniformly over cell, twice as numerous as long spine scales, to 4-6 um long, structured like
long spine scales but the supporting struts greatly reduced, base-plate elliptical, c. 1.5 * 2
um; plate scales forming an underlayer, flat, elliptical, c. 1.3 * 1.6 um, apparently rimless,
patterned with delicate ±radiating striae. Flagella equal, to 20 um long. Haptonema
somewhat longer than flagella when extended, or coiled into c. 10 gyres when retracted.
distribution: Southern Ocean, south of Australia; Cape Town, South Africa (Manton,
1978); Arctic waters (Manton, 1978).

The haptonema of C. hirta has been shown to be a food-capturing device (Kawachi et al.,
1991). This species can ingest particulate matter of size range 0.5-1.97 um diameter
(Marchant, unpublished data).

Chrysochromulina leadbeateri Estep, in Estep et al. (1984: 618, figs 6, 7)

ADDITIONAL REFERENCES: Leadbeater (1972b: figs 3-5), Hallegraeff (1983: 498, fig. 16, as
Chrysochromulina sp. 1), LeRoi & Hallegraeff (2004: 83, figs 18, 19)

Fig. 5.4b

description: Cells spherical, 1.5-4 um diameter (dried material, possibly up to twice as

large when living), biflagellate; body covered with unmineralised body scales. Body scales
of 2 types: both types circular, 0.2-0.4 um diameter, with a cruciform central pattern
surrounded by a 0.03 um diameter central ring; one scale type with an additional raised ring
near periphery and a radial pattern of 25-30 striae on one face; other scale type with a
slightly raised rim and little or no patterning. Flagella slightly subequal, c. 11 and 13 um
long. Haptonema longer than flagella when extended, in 8-18 gyres when retracted.

distribution: Southern Ocean, south of Australia; North Atlantic (Estep et al, 1984);
eastern Australia (Hallegraeff, 1983); Norway (Leadbeater, 1972b).

Chrysochromulina polylepis Manton & Parke (1962: 575, figs 1-28)

ADDITIONAL REFERENCES: Johnsen et al. (1992: 241), Edvarsen & Vaulot (1996: 94), Throndsen
(1997: 642, pi. 9), LeRoi & Hallegraeff (2004: 87, fig. 27)

Fig. 5.5d, e

description: Cells ovoid to subspherical, with the flagellar pole obliquely truncate and
depressed centrally, 6-12 um long, 5-9 um wide, biflagellate; body covered in unmineral-
ised scales. Body scales of 4 types: large plates with upturned rims, elliptical, c. 1.4 x 1.2
um; numerous small plates with rims flexed towards distal surface and with 2 central
bulbosities on distal surface, elliptical, c. 1.2 x 0.8 um; narrow elliptical plates with the rim
flattened against distal face, c. 1.2 x 0.6 um, probably present in a group near flagellar pole;
a few large elliptical plates, c. 2.5 * 0.9 um, with rims and a forked excresence at one end,
situated at the flagellar pole. Flagella equal or subequal, arising from a central depression,
c. 25 um long, 2-3 times length of cell body. Haptonema c. 15 um long, 1-1.5 times body

length when extended.

distribution: Southern Ocean, south of Australia; northern seas at 50°11'N 04°13'W

(Manton & Parke, 1962); coastal, oceanic Europe (Throndsen, 1997).

Chrysochromulina vexillifera Manton & Oates (1983a: 519, figs 2-16; type fig. 5c)

Fig. 5.7a-d

description: Cells ovoid, c. 8 * 6 um, biflagellate; body covered in unmineralised scales.

Body scales of 3 types: 14 large spinous scales at each end of cell, with spine to 15 um long,
tapered, pointed and supported basally by 4 struts attached to base-plate, base-plate strongly

262
 HAPTOPHYTES: ORDER PRYMNESIALES

\\

-A

Figure 5.5. Chrysochromulina species, a-c, C. ericina. a, TEM Cell with flagella, but most
scales have been dislodged from the cell body, b, TEM Cluster of dislodged spine scales and
plate scales, c, TEM Detail of long spine scale and circular plate scale, d, e, C. polylepis. d,
TEM One small and three large body plate scales, e, TEM Proximal surface of narrow elliptical
plate scale from flagellar pole area. Scale bars: a-c = 2 urn; d, e = 0.5 um.

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 HAPTOPHYTES: ORDER PRYMNESIALES

Figure 5.6. Chrysochromulina hirta. a, TEM Collapsed cell, with flagella. b, TEM Flagella
and plate scales, c, TEM Detail of long (lsp) and short (ssp) spine scales, d. TEM Detail of
plate scales. Scale bars: a-c = 2 urn; d =
1 urn.

264
 HAPTOPHYTES: ORDER PRYMNESIALES

Figure 5.7. Chrysochromulina vexillifera. a, TEM Large (lpl) and small (spl) plate scales, b,
TEM Spine scale (sp) and large plate scales, c, TEM Detail of spine scale and large and
small (spl) plate scales, d, TEM Detail of outer (os) and inner (is) surfaces of large plate
scales. Scale bars: a-d = 2 \im.

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 HAPTOPHYTES: ORDER PRYMNESIALES

Figure 5.8. Chrysochromulina types A and B. a, b, type A. a, TEM Collapsed cell with
dislodged scales, b, TEM Four scale types: long (lsp) and short (ssp) spine scales, and large
(Ipl) and small (spl) plate scales, c, d, type B. c, TEM Striated surfaces of large and small
plate scale, d, TEM Fibrous surface and partly collapsed rim of large plate scale. Scale bars:
a = 2 urn; b-d = 1 um.

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 HAPTOPHYTES: ORDER PRYMNESIALES

convex; large outer plate scales, flat, circular to elliptical, 1.6-2 urn diameter, with a narrow
thickened margin, distal surface with a fibrous pattern and bearing 4 vestigial struts, with
spine absent, proximal surface patterned with c. 130 fine radiating striae; underlayer of small
elliptical plate scales, 0.5 * 0.8 urn, with a narrow raised rim and faint patterning of radial
striae on proximal face, patternless on distal face. Flagella c. 18-20 um long. Haptonema at
least 13 um long when extended.

distribution: Southern Ocean, 45.38-52.01 °S 134. 05-144. 20°E, south of Australia;

Galapagos Is. (Manton & Oates, 1983a).

Chrysochromulina type A
Fig. 5.8a, b

description: Cell body 4.3-5 \im diameter; flagella and haptonema not observed. Body scales
of 4 types: a small number of long spine scales with an upwardly flexed rim, with the spine
supported by 4 struts, overall length 10-20 jam; short spine scales with an upwardly flexed rim,
distal surface of base-plate with c. 80 fine radiating striae, with the spine supported by 4 struts,
overall length 0.3 urn; large elliptical plate scales 1 * 1.3 um, distal surface with c. 80 fine
radiating striae and a slightly thickened rim, proximal surface with a fibrous pattern; small
elliptical plate scales, 0.5 * 0.7 um, finely striated on one surface, patternless on the other.

distribution: Southern Ocean, south of Australia, 45. 38-57. 31°S 101.18-144.20°E.

Our material closely resembles both C. vexillifera and our Chrysochromulina type C. Type A
differs from having 4 types of scales (3 types in vexillifera), as well as some
C. vexillifera in
fine details of scale morphology. Type A differs from type C in the fine detail of scale
morphology of the short spine scales. Until further material becomes available, we regard the
3 collections as representing different species, despite comparatively minor morphological
differences.

Chrysochromulina type B
Fig. 5.8c, d

description: [Only scales observed.] Body scales of 2 types: large plate scales, elliptical, 1.3 *
1.9 um, with a delicate raised margin, proximal surface with a 0.18 um wide rim and c. 140—
150 fine striae, distal surface with a fibrous pattern; small plate scales elliptical, 0.7 * 1 um,
one surface with c. 60 fine radiating striae.

distribution: Southern Ocean, south of Australia, 45. 38-57. 31°S 77. 35-144. 20°E.

The scales are similar to those of C. chiton, except that in type B the rims of the larger scales
are not striated, as they are in C. chiton.

Chrysochromulina type C
Fig. 5.9a-d

description: [Only scales observed.] Body scales of 4 types: large spine scales, with the
spine tapered, supported basally by 4 struts 1.4 um long, attached to base-plate, overall
length to 15 um, base-plate upwardly flexed, distal surface with fine radiating striae; small
spine scales similar in structure to large spine scales, but overall length 0.6-1.6 um, distal
surface with 80-100 fine radiating striae; large outer plate scales, flat, circular to elliptical,
c. 2 um diameter, with a faint rim, distal surface patterned with c 110 fine radiating striae;

underlayer of small elliptical plate scales 0.5 * 0.8 um, with a narrow raised rim, proximal
surface with a faint patterning of radial striae, distal surface patternless.

distribution: Southern Ocean, Pacific sector, 54°30.4'S 158°47.0'E.

Our material closely resembles C vexillifera. See discussion under Chrysochromulina type A.

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 HAPTOPHYTES: ORDER PRYMNESIALES

Chrysochromulina type D
Fig. 5.10a, b

description: Cell body spherical, 2.3-4.6 um diameter. Body scales of 1 type: plate scales
elliptical, 0.6 * 0.8 um;, distal surface possibly concave, with c. 16 rather coarse radiating
striae and a thickened margin.

distribution: Southern Ocean, south of Australia, 54.02-57.3 1°S 120.04-1 28. 17°E.

Chrysochromulina type E
Fig. 5.10c

description: [Only scales observed.] Body scales of 2 types: short spine scales with base-
plate elliptical, 1.2 * 1.6um, distal surface with a fibrous pattern and 4 ridges supporting a
central c. 0.25 urn long spine; plate scales elliptical, 0.8 * 1.1 um, with c. 76 fine radiating
striae on one surface, other surface with a fibrous pattern.

distribution: Southern Ocean, south of Australia, 46.20-57.3 1°S 101.18-134.05°E.

Chrysochromulina type F
Fig. 5.10d

description: [Only scales observed.] Body scales of 1 type: spine scales with base-plate
circular to oval, 2-2.4 um diameter, with an upwardly flexed rim and a cross-hatched fibrous
pattern on distal surface; spine supported by 4 struts; overall length 3.5 um.

distribution: Southern Ocean, south of Australia, 52.01 °S 134.05°E.

Chrysochromulina type G
Fig. 5.11a, b

description: [Only scales observed.] Body scales of 1 type: spine scales with a base-plate
circular to elliptical, 2-2.4 um diameter; proximal surface with fine striae; spine supported
by 4 arched struts; overall length of spine and struts 0.5 um.
distribution: Southern Ocean, south of Australia, 46.20-49.47°S 85.1 7-1 01.18°E.

Chrysochromulina type H
Fig. 5.11c, d

description: [Only one cell observed.] Cell spherical, 2 um diameter. Flagella c. 12 um

long. Body scales of 1 type: plate scales sparse, circular, 0.5-0.7 um diameter, with 12-14
fine radiating striae on one (?distal) surface.

distribution: Southern Ocean, south of Australia, 52.0 1°S 134.05°E.

The body scales of the single specimen collected resemble C. mantoniae B.Leadb. from
Tasmania (LeRoi & Hallegraeff, 2004), but the haptonema is considerably longer (Fig. 5.1 lc)
than the 10 um reported for C. mantoniae. The latter species also bears long spine scales at
the anterior and posterior ends of the obovate cell body.

Chrysochromulina type I

Fig. 5.12a-d

description: [Only one cell observed.] Cell body 3.7 um diameter; flagella and haptonema
not seen. Body scales of 4 types: large spine scale base circular, c. 1 um diameter, with an
upwardly flexed rim, spine 10 um long overall; small spine scale base elliptical, 0.4 * 0.4 um,
with a distinct rim, spine 0.35 um long; large plate scale elliptical to circular, 1.1 * 1.6 um,

268
 HAPTOPHYTES: ORDER PRYMNESIALES

Figure 5.9. Chrysochromulina type C. a, TEM Long and short spine scales, b, TEM
Dislodged small spine scales, c, TEM Small spine scales and small (spl) and large (Ipl) plate
scales, d, TEM Detail of small spine scales. Scale bars: a-d =1um.

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 HAPTOPHYTES: ORDER PRYMNESIALES

Figure 5.10. Chrysochromulina types D, E and F. a, b, type D. a, TEM Scale mass dislodged
from cell body, b, TEM Detail of scales, c, type E, TEM Spine scales and small plate scales, d,
type F, TEM Spine scales with distinct, cross-hatched surface pattern. Scale bars: a-d =1urn.

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 HAPTOPHYTES: ORDER PRYMNESIALES

Figure 5.11. Chrysochromulina types G and H. a, b, type G, TEM Spine scales, c, d, type H.
c,TEM Cell with flagella and haptonema. d, TEM Plate scales with sparse surface striations.
Scale bars: a, b, d =
1 urn; c = 2 urn.

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 HAPTOPHYTES: ORDER PRYMNESIALES

Figure 5.12. Chrysochromulina type I. a, TEM Whole cell with dislodged spine scales and
plate scales, b, TEM Long spine scale, c, TEM Detail of base of long spine scale and both
surfaces of large plate scales, d, TEM Distal (os) and proximal (is) surfaces of small plate
scales, and small spine scale (ssp). Scale bars: a = 2 urn; b-d = 1 pm.

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 HAPTOPHYTES: ORDER PRYMNESIALES

Figure 5.13. Chrysochromulina types J and K. a, b, type J. a, TEM Collapsed cell with
dislodged scales, b, TEM Detail of striated plate scales, c, d, type K. c, TEM Whole cell with
attached body scales, d, TEM Detail of plate scales. Scale bars: a = 1 pm; b-d = 0.5 pm.

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 HAPTOPHYTES: ORDER PRYMNESIALES

Figure 5.14. Chrysochromulina types L and M. a, b, type L. a, TEM Whole cell with
flagella and dislodged scales, b, TEM Detail of striated surface of scales, c, d, type M. c,
TEM Whole cell with flagella and coiled haptonema. d, TEM Detail of scale surfaces. Scale
bars: a-d = 1 urn.

274
 HAPTOPHYTES: ORDER PRYMNESIALES

distal surface with c. 90 proximal surface with an interwoven fibrous

fine radiating striae,
c. 68
pattern; small plate scale elliptical to circular, 0.4 * 0.5 urn, distal (outer) surface with
fine radiating striae and 2 off-centre holes, proximal (inner) surface with a distinct rim, an
interwoven fibrous pattern and 2 off-centre holes.

distribution: Southern Ocean, south of Australia, 46°24.7'S 139°21.6'E.

Chrysochromulina type J
Fig. 5.13a, b

description: [Only one cell observed.] Cell body 2.8 pm diameter. Body scales of 1 type:
plate scales circular, 0.6 um diameter, with a thickened rim and c. 24 radiating striae.

distribution: Southern Ocean, south of Australia, 52.01°S 134.05°E.

Chrysochromulina type K
Fig. 5.13c, d

description: [Only one cell observed.] Cell body 2.8 um diameter. Body scales of 1 type:
short spine scales elliptical to circular, 0.2 * 0.3 um; distal surface with very faint striae;
spine supported by 3 ridges (the shadow casting preparation often makes it resemble a cross
or the numeral '4'); spine 0.15-0.2 um long.

distribution: Southern Ocean, offshore Prydz Bay, East Antarctica, 64°14.4'S 79°07.8'E.

Chrysochromulina type L
Fig. 5.14a, b

description: [Only one cell observed.] Cell body spherical, 2.8 um diameter. Flagella
unequal, to 14 pm long. Body scales of 1 type: plate scales subcircular, 0.8-0.9 pm diameter;
one surface with c. 30 radiating striae, each distinctly bifurcated near the scale margin; other
surface patternless.

distribution: Southern Ocean, south of Australia.

Chrysochromulina type M
Fig. 5.14c, d

DESCRIPTION: [Only one cell observed.] Cell body ovoid, 2.4 pm long, 1.7 pm diameter.
Flagella equal, to 4.2 pm long; haptonema c. 10 pm long. Body scales of type: plate scales
1

subcircular, 0.8-0.9 pm diameter; one surface with c. 30 radiating striae, each distinctly
bifurcated near the scale margin; other surface patternless.

DISTRIBUTION: Southern Ocean, south of Australia.

275
 HAPTOPHYTES:
6.

ORDER COCCOLITHOPHORALES*
Claire S. Findlay, Jeremy R. Young and Fiona J. Scott

Coccolithophores are single-celled marine algae + which are distinguished from other
phytoplankton by an external covering of calcite scales, or coccoliths. They are confined to
the photic zone (the uppermost 150-200 m) of the oceans where they are one of the principal
primary producers and a significant contributor to biomass. Reproduction rates are estimated
at up to 2.5 divisions per day (Brand, 1994), with a turn-over of 4 to 10 days (Honjo, 1976)
and standing stocks of up to 10 million specimens per litre (Winter et al., 1994). Their bloom
phases are clearly visible in satellite images, turning the sea white (Groom & Holligan, 1987;
Holligan et al., 1993; Tyrell et al, 2004).

The coccolithorphore life cycle typically consists of an alternation of a diploid, motile or

non-motile phase, usually bearing heterococcoliths (calcite scales constructed of complex
calcite crystals of various shapes and sizes) and a haploid, predominantly motile phase,
which, if calcified, produces holococcoliths (calcite scales constructed of small calcite
crystals of uniform size and shape). This type of life cycle was first documented by Parke &
Adams (1960) for Coccolithus pelagicus (G.C.Wall.) J.Schiller. Billard (1994) synthesised a
range of data to predict that it was a common pattern, and this has now been confirmed by
Houdan et al. (2004). Unusually, both life cycle phases are capable of indefinite asexual
reproduction, and this seems to be a key adaptation allowing the group to flourish in
heterogeneous environments.

Motile coccolithophores typically bear two smooth, equal flagella and a haptonema (Billard,
1994). The haptonema is a unique feature of haptophyte planktonic algae; it is similar to a
flagellum, but it has a different microtubular ultrastructure. This structure bends, rotates,
coils and attaches to substrata and has been observed capturing particles (Inouye & Kawachi,
1994).

Coccospheres can have non-calcified scales in addition to coccoliths. In some species they
have two or more layers of the same type of coccolith, e.g. Calcidiscus leptoporus, Emiliania
huxleyi. Other taxa develop dithecate coccospheres with two layers of different types of
coccoliths, an inner endothecal layer and an outer exothecal layer, e.g. Syracosphaera
pulchra.

The taxonomy of coccolithophores is primarily based on the structure and crystallography of

the coccoliths. Detailed descriptions of coccolith and calcareous nanofossil terminology can
be found in Kleijne (1993), Winter & Siesser (1994), Jordan et al. (1995) and Young et al.
(1997). For more detailed information on the biology, morphology, ecology and
biogeography of coccolithophores, refer to Winter & Siesser (1994), Green & Leadbeater
(1994) and Thiersten & Young (2004). The taxonomy of coccolithophores has been reviewed
by Cros & Fortuno (2002) and Young et al. (2003).

This exceptional group of phytoplankton is a significant component of biogeochemical

pathways, e.g. the global carbon-carbonate cycle, operating as a major sink of carbon dioxide
and source of carbonate in the oceans (Honjo, 1976; Westbroek et al., 1994). Through
deposition on the seafloor, coccolithophores have a profound effect on the deep sea carbon
dioxide budget, calcite compensation depth, lysocline and carbonate turnover.

As producers of dimethylsulphoniopronate (DMSP) and dimethylsulphide (DMS), these

organisms are an important component of global sulphur cycles (Holligan et al., 1987;
Turner et al., 1988; Malin et al., 2004). DMS is released into the atmosphere and is

Order Coccolithophorales of Class Prymnesiophyceae.

+ There is one extant freshwater species, Hymenomonas roseola Stein.

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 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

converted by oxidation to bi-products that act as nuclei for cloud formation, increasing cloud
albedo, or reflectivity, a significant component in climate variation (Gibson et al., 1990;

Malin et al., 1994; Malin & Kirst, 1997).

From a geological perspective, the carbonate remains of these phytoplankton are the main
component deep sea calcareous ooze and chalk deposits, covering approximately one-third
in
of the Earth's surface. They first appear in the fossil record in the Late Triassic, abundant
with low diversity and restricted to low latitudes with only one species surviving the
Triassic/Jurassic extinction event (Bown et al., 2004). They rapidly re-established in all

marine environments in the Early Jurassic and progressively diversified through the
Mesozoic, reaching a diversity peak in the Late Cretaceous. Subsequently, approximately
90 percent of species disappeared during the Cretaceous/Tertiary extinction event (Bown &
Young, 1998). The group rapidly recovered with a second diversity peak in the Mid-Eocene.
As these nanofossils are abundant, evolve rapidly and are cosmopolitan, they are an excellent
biostratigraphic tool, especially for global correlation of oceanic sediments.

In the fields of palaeoceanography and palaeoclimatology, studies of calcareous nanofossils

offer an excellent analytical tool. Various coccolithophore assemblages are associated with
specific water masses reflecting the physical and chemical properties of those surface waters.
These assemblages, preserved in the deep-sea sediments, provide a proxy for interpreting
changes of temperature, salinity, nutrient levels, turbidity and ocean circulation within
surface waters through time. They have been used effectively to interpret changes in the
position of oceanic fronts in the Southern Ocean (Wells & Connell, 1997; Findlay, 1998;
Findlay & Flores, 2000; Findlay & Giraudeau, 2002), the sub-Antarctic Pacific Ocean
(Geitzenauer, 1969), the Nordic Seas (Andruleit & Baumann, 1998; Baumann &
Mattheissen, 1992) and the Mediterranean (Flores et al, 1997) and changes in oceanic
currents south of Africa (Flores et al, 1999). Preserved assemblages have been used to
monitor carbonate dissolution over time (Berger, 1973a, 1973b). Some species of
coccolithophores are associated with long-chain alkenones constituting up to 8 percent of
their biomass. The unsaturation in these alkenones is a function of temperature, providing a
further tool for palaeoceanography and palaeoclimatology research (Prahl & Wakeham,
1987; Prahl et al, 1988; Marlowe et al., 1990; Baumann et al, in press).

The biogeography of living calcareous phytoplankton in the high latitudes of the Southern
Ocean has only been studied in comparatively recent times (Nishida, 1986; Findlay, 1998;
Eynaud et al, 1999; Findlay & Giraudeau, 2000). The results document a decrease of
abundance and diversity of coccolithophores in a poleward direction in the Southern Ocean
in conjunction with decreasing water temperature and salinity and an increase in levels of
nitrateand phosphate, with highest abundance and diversity recorded across the Subtropical
Front and Sub-Antarctic Front. Separate assemblages, all dominated by Emiliania huxleyi
(Type B/C, which appears to be unique to the Southern Ocean), are associated with distinct
surface and subsurface water masses. Two assemblages are recognised in the polar region of
the Southern Ocean, a monospecific assemblage of Emiliania huxleyi at 52-60°S, and a
semi-calcified assemblage poleward of the Antarctic Divergence (63°S).

Thirteen families of calcareous nanoplankton are recognised in Order Coccolithophorales, and

six have been identified in the polar regionof the Southern Ocean, i.e. south of the Polar
Front (as defined by Belkin & Gordon, 1996). Eleven genera and 20 species have been
positively identified in this region (Table 6.1). Specimens that have not been unequivocally
identified are not included in this review, e.g. "holococcolithophore spp." (Eynaud et al,
1999) and "Calciopappus sp." (Nishida, 1986). A subtropical assemblage identified by Winter
et al (1999) in the Weddell Sea is excluded as it is probably either the result of contamination
or may represent a "captured" assemblage transported south in a pocket of warm, subtropical
water. In the Weddell Sea, coccospheres were identified by SEM in two samples in the polar
region, one collected by a plankton tow net at depths of 0-50 m [Calcidiscus leptoporus
(1 coccosphere) and Emiliania huxleyi (2)], and the other via the ship's pump from 8 m water
depth [E. huxleyi (34), Discosphaera tubifera (1), Gephyrocapsa muellerae (4), Syraco-
sphaera pulchra (1), S. anthos (1) and Umbilicosphaera hulburtiana (4)]. Isolated coccoliths
are also excluded from this review, as they do not accurately represent living assemblages.

277
 -

HAPTOPHYTES: ORDER COCCOLITHOPHORALES

Table 6.1. Coccolithophores identified from the Southern Ocean south of the Polar Front.

Species Latitude Longitude Reference

Calcidiscus leptoporus HET 57°31'S 150°02'E Nishida(1986)

Calcidiscus leptoporus HOL 64°16'S 136°06'E Nishida(1986)
65°00'S 117°59'E Nishida(1986)
60°00'S 116°02'E Nishida(1986)
57°29'S 115°59'E Nishida(1986)
53°00'S 06°irw Eyaund et al. (1999)
Helicosphaera carter i HET 57°31'S 150°02'E Nishida(1986)
Helicosphaera carteri HOL 64°16'S 136°06'E Nishida(1986)
60°00'S 116°02'E Nishida(1986)
Emilia nia huxleyi 57°31'S 150°02'E Nishida(1986)
61°24'S 150°05'E Nishida(1986)
64°56'S 150°12'E Nishida(1986)
64°16'S 136°06'E Nishida(1986)
65°00'S 117°59'E Nishida(1986)
60<WS 116°02'E Nishida(1986)
57°29'S 115°59'E Nishida(1986)
63°17'S 90°00'W Nishida(1986)
64 18'S 90°00'W Hasle(1969)
62°01'S 120°00'W Hasle(1969)
63°03'S 120°00'W Hasle(1969)
50°25'S 00°47'W Eyaund et al. (1999)
50°45'S 01°24'W Eyaund et al. (1999)
51°10'S 02°06'W Eyaund et al. (1999)
51°30'S 02°43'W Eyaund et al. (1999)
52°00'S 03°36'W Eyaund et al. (1999)
52°30'S 04°47'W Eyaund et al. (1999)
53°00'S 06°11'W Eyaund et al. (1999)
53°59'S 09°04'W Eyaund et al. (1999)
55°00'S 12°06'W Eyaund et al. (1999)
56°01'S 15°12
-

W Eyaund et al. (1999)

57°01'S 18°15'W Eyaund et al. (1999)
55°01'S 141°00'E Findlay & Giraudeau (2000)
55°55'S 140°24'E Findlay & Giraudeau (2000)
57°23'S 139°50'E Findlay & Giraudeau (2000)
58°51'S 139°50'E Findlay & Giraudeau (2000)
60°21'S 139°50'E Findlay & Giraudeau (2000)
61°20'S 139°50'E Findlay & Giraudeau (2000)
63°21'S 139°50'E Findlay & Giraudeau (2000)
Gephyrocapsa oceanica 64°56'S 150°12'E Nishida(1986)
65°00'S 117°59'E Nishida(1986)
Gephyrocapsa caribbeanica 64°16'S 136°06'E Nishida(1986)
Gephyrocapsa muellerael See G. caribbeanica
Gephyrocapsa ericsonii 64°16'S 136°06'E Nishida(1986)
65°00'S 117°59'E Nishida(1986)
Pappomonas weddellenis 65°39'S 47°34'W Thomsen et al. (1988)
64°49'S 44041 w Thomsen et al.(1988)
Papposphaera obpyramidalis 61°20'S 139°50'E Findlay & Giraudeau (2000)
64°49'S 43°42'W Thomsen et al. (1988)
65°26'S 45°38'W Thomsen et al. (1988)
65°39'S 47°34'W Thomsen et al. (1988)
64°55'S 48°04'W Thomsen et al. (1988)
Papposphaera boreal is HET 58°51'S I39°50'E Findlay & Giraudeau (2000)

278
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

Table 6.1. Continued.

Species Latitude Longitude Reference

Papposphaera borealis HOL 61 2(TS 139°50'E Findlay & Giraudeau (2000)

Papposphaera arctica HOL 64°51'S 46°05'W Thomsen el al. (1988)
Papposphaera polybotrys HOL Thomsen et al. (1991a)
Papposphaera simplicissima 64°49'S 4404I 'W Thomsen e! al. (1988)
64°49'S 44-41 'W Thomsen et al. (1988)
65°00'S 49°54'W Thomsen et al. (1988)
64°51'S 45°55'W Thomsen et al. (1988)
Syracosphaera pulchra HET 64°16'S 136°06'E Nishida(1986)
Syracosphaera pulchra HOL 60°00'S 116°02'E Nishida(1986)
Syracosphaera molischii HET 55°00'S 141°10'E Van den Enden, unpubl.
Umbellosphaera irregularis 57°31'S 150°02'E Nishida(1986)
64°56'S 150°12'E Nishida (1986)
Wigwamma antarctica 6P20'S 139°50'E Findlay & Giraudeau (2000)
64°49'S 43°42'W Thomsen et al. (1988)
65°39'S 47°34'W Thomsen et al. (1988)
65°33'S 47°40'W Thomsen et al. (1988)
64°49'S 4404I 'W Thomsen et al. (1988)
64°51'S 46°05'W Thomsen et al. (1988)
65°09'S 43°03'W Thomsen et al. (1988)
Wigwamma arctica 64°39'S 43°36'W Thomsen et al. (1988)
64°49'S 44 41'W Thomsen et al. (1988)
Wigwamma annul ifera HET 64°5LS 46°05'W Thomsen et al. (1988)
Wigwamma annulifera HOL 65°09'S 43°03'W Thomsen et al. (1988)
Wigwamma triradiata 61°20'S 139°50'E Findlay & Giraudeau (2000)
64°49'S 43°42'W Thomsen et al. (1988)
64°51'S 46°05'W Thomsen et al. (1988)
Trigonaspis melvillea 64°49'S 43°42'W Thomsen et al. (1988)
65°26'S 45°38 ,

W Thomsen et al. (1988)

65°39'S 47°34'W Thomsen et al. (1988)
65°16'S 44°12'W Thomsen et al. (1988)
64°49'S 44°41'W Thomsen et al. (1988)
64°51'S 46°05'W Thomsen et al. (1988)

The total diversity of coccolithophores recorded of the Southern Ocean

in the polar regions
is significantly lower than in the sub-Arctic (as reviewed by Samtleben et al., 1995);
abundance is also typically lower. In the subpolar North Atlantic, coccolithophores,
especially Coccolithus pelagicus, often dominate assemblages. Conversely, in the polar
regions of the Southern Ocean, coccolithophores are almost always subordinate to diatoms.
This is probably a reflection of the major biogeochemical differences between the sub-Arctic
and the polar regions of the Southern Ocean where iron limitation appears to be a major
controlling factor of total productivity. In the Southern Ocean iron inputs are extremely low
and, consequently, macronutrients, including silica, are rarely reduced to trace levels, and
siliceous diatoms usually dominate phytoplankton assemblages. By contrast, iron is rarely
limiting in the subpolar North Atlantic; under these conditions coccolithophores can replace
diatoms as the dominant group when silica becomes limiting.

The is that of Kleijne (1993), Jordan & Kleijne (1994) and Young
classification followed here
et al. (2003).This latest synthesis (Young et al, 2003) provides an overview of nanoplankton
taxonomy and a guide to the identification of coccolithophores. The descriptions are taken
partly or completely from Young et al. (2003), supplemented by Jordan & Kleijne (1994).
More detailed descriptions can be found in these publications, the original publications for
each species and Kleijne (1993). For higher classification, the key diagnostic criteria are often

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 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

crystallographic orientation, especially the relative development of cycles of elements with

subvertical crystallographic orientation (V-units) and those with subradial orientation (R-
units).This aspect is not covered in this treatment since it requires specialist study and is not
necessary for basic identification. For details see Young et al. (2003).

Quoted coccolith sizes are based on the long axis of the distal shield. Recent research has
shown that the typical coccolithophore has numerous life cycle associations of holococco-
lithophore and heterococcolithophore species. Following Thomsen et al. (1991b), Cros et al.
(2000) and Young et al. (2003), a single species name is applied to both phases of the life
cycle with an informal indication of which phase is observed, i.e. HET (heterococco-
lithophore) and HOL (holococcolithophore).

Heterococcolithophores

Cells covered in heterococcoliths, i.e. coccoliths formed of complex crystals arranged in

radial arrays.

Family Calcidiscaceae
Diploid phase non-motile, with coccospheres formed of robust placoliths. Haploid phase
motile, with holococcoliths.

Calcidiscus

Coccoliths (sub-)circular; central area closed or with a narrow opening; distal shield with
laevogyral sutures.

Calcidiscus leptoporus (G.Murray & V.H.Blackman) Loebl. & Tappan (Loeblich &
Tappan, 1978: 1391, not illustrated) HET
BASIONYM: Coccosphaera leptopora G.Murray & V.H.Blackman (1898: 430, pi. 15, figs 1-7)

Figs 6.1a, c; 6.2a

description: Three morphotypes (large, intermediate and small) have been recognised
(Kleijne, 1993; Knappertsbusch et al., 1997) and shown to represent discrete taxa (Geisen et
al, 2002):

Calcidiscus leptoporus subsp. leptoporus HET

Intermediate morphotype (Fig. 6.1c): coccoliths 5-8 urn; distal shield with 15-30 elements
and smoothly curved sutures. The most common form.

HOL phase synonym: Crystallolithus rigidus Gaarder, in Heimdal & Gaarder (1980)

Details of holococcolithophores are given separately to avoid confusion (p. 292).

Calcidiscus leptoporus subsp. quadriperforatus (Kamptner) Geisen et al. (2002: 543,

figs 18, 20-22) HET
BASIONYM: Syracosphaera quadriperforata Kamptner (1937: 302, pi. 15, pp. 15-16)

Large morphotype (Fig. 6.1a): coccoliths 7-11 um; distal shield with 20-35 elements and
smoothly curved sutures.

HOL phase synonyms: Syracosphaera (Syracolithus) quadriperforata Kamptner (1941)

Syracolithus quadriperforatus (Kamptner) Gaarder (1962)

280
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

Calcidiscus leptoporus HET

Small morphotype (Fig. 6.2a): coccoliths 3-5 um; distal shield with 10-20 elements, usually
with angular or serrated sutures (Kleijne, 1993).

Family Helicosphaeraceae
Diploid phase motile; coccospheres ellipsoidal, with a distinct flagellar opening; coccoliths
(helicoliths) arranged spirally around coccosphere, varying in size, larger and with more
prominent wings toward the flagellar end. Coccoliths with a helical flange terminating in a
wing. Haploid phase motile, with holococcoliths formed of crystallites arranged in
rhombohedral arrays.

Helicosphaera
Coccoliths with a helical flange; central area forming a slit, with or without a disjunct bar.

Helicosphaera carteri (G.C.Wall.) Kamptner (1954: 21, 23, figs 17-19) var. carteri HET
BASIONYM: Coccosphaera carteri G.C.Wall. (Wallich, 1877: 348, pi. 17, figs 3, 4, 6, 7, 17), as carterii

Figure 6.1. Calcidiscus leptoporus. a, C. leptoporus subsp. quadriperforatus (HET), large

morphotype, SEM. b, C. leptoporus subsp. quadriperforatus (HOL), SEM (courtesy of
J.Bollman, EMIDAS database), c, C. leptoporus subsp. leptoporus (HET), intermediate
morphotype, SEM. d, C. leptoporus subsp. leptoporus (HOL), SEM (courtesy of S.Renaud,
EMIDAS database). Scale bars: a = 1 urn; b-d = 2 |im.

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 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

SYNONYMS: Helicopontosphaera kamptneri W.W.Hay & H.Mohler, in Hay et al. (1967)

Helicosphaera kamptneri (W.W.Hay & H.Mohler) Locker (1973)

Fig. 6.2b

DESCRIPTION: Medium to large coccoliths 6-12 urn; flange terminating in wing; central area
of 2 in-line slits or a single long slit.

HOL phase synonyms: Syracolithus catilliferus (Kamptner, 1937)

Syracolithus confusus Kleijne (1991)

Both of these holococcolithophore species have been found in combination form with H.
carteri.

Figure 6.2. Calcidiscus leptoporus and Helicosphaera carteri. a. C. leptoporus. small

morphotype, SEM. b, H. carteri (HET), SEM. c, H. carteri (HOL), solid variant form, SEM
(courtesy of S.Renaud, EMIDAS database), d. //. carteri (HOL), perforated variant form,
SEM (courtesy of M.Knappertsbusch, EMIDAS database). Scale bars: a-d = 2 um.

282
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

Family Noelaerhabdaceae
Diploid phase a non-motile coccosphere; overlapping placoliths with a reticulum or grill
covering the central area opening. Haploid phase motile and non-calcifying.

Emiliania

This genus is characterised by slits in the distal shield, sometimes referred to as "T" elements.

Emiliania huxleyi (Lohmann) W.W.Hay & H.Mohler, in Hay et al. (1967: 447, pi. 10-12,
figs 1,2)

BASIONYM: Pontosphaera huxleyi Lohmann (1902: 129, 130, pi. 4, figs 1-9; pi. 6, fig. 6a)

Fig. 6.3a

Figure 6.3. Emiliania huxleyi and Gephyrocapsa species, a, E. huxleyi, Type B/C, SEM. b,
G. oceanica, SEM. c, G. muellerae, SEM. d, G. ericsonii, SEM (courtesy of J.Bollman,
EMIDAS database). Scale bars: a-c = 2 um; d = urn. 1

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 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

description: Three morphotypes were recognised by Young & Westbroek (1991), and an
additional morphotype appears to be present in the Southern Ocean:

Type A (huxleyi): Coccoliths 3^4 um; distal shield elements robust; central opening covered
by a grill.

Type B (pujosae): Coccoliths 3.5-5 um; distal shield elements delicate; central area covered
with lath-like elements.

Type C (kleijneae): Coccoliths 2.5-3.5 um; distal shield elements delicate; central area open
or covered by a thin plate.

Type B/C: Coccoliths 3-4 um; elements delicate; central area open or covered
distal shield
by a thin plate. This is similar in to Types B and C but is intermediate in size.
morphology
This form dominates assemblages in the Southern Ocean (Hockfield, 2000; Findlay &
Giraudeau, 2000).

Gephyrocapsa
Coccoliths with the central area spanned by a bridge structure; proximal and distal shields
usually solid.

Gephyrocapsa caribbeanica Boudreaux & W.W.Hay, in Hay et al. (1967)

description: Coccoliths of intermediate size, with a low to intermediate bridge angle;

central area closed or almost closed.

Size and bridge angle of G. caribbeanica are very similar to those of G. muellerae, the only
apparent difference being a closed central area in G. caribbeanica. The latter is rarely found
in living assemblages but is common in Late Quaternary sediments. Records of
G. caribbeanica from the plankton in older literature (e.g. Nishida, 1986) mostly refer to
morphotypes that would now be placed in G. muellerae. Bollman (1997) provided a
summary of Gephyrocapsa species and included the measurements for G. caribbeanica
(holotype) with a bridge angle of 45° and coccolith length of 4.1 um (Boudreaux & Hay,
1969). Gephyrocapsa muellerae (holotype) has an bridge angle of 23° and coccolith length
of 3.9 um (Breheret, 1978).

Gephyrocapsa ericsonii Mclntyre & Be (1967: 571, pi. 10; pi. 12, fig. b)

Fig. 6.3d

description: Coccospheres 3-5 um. Coccoliths very small, 1.4-2.3 um, with a low to
intermediate bridge angle; central area wide; bridge thin.

Gephyrocapsa muellerae Breheret (1978: 448, pi. 2, figs 3, 4)

Fig. 6.3c

description: Coccospheres 5-9 um. Coccoliths of intermediate size, 3-4 um, with a low to
intermediate bridge angle; central area small.

Gephyrocapsa oceanica Kamptner (1943: 45)

Fig. 6.3b

description: Coccospheres 6-10 um, often in clusters. Coccoliths large, 3.5-6 um, with
bridge at a high angle to the long axis; collar around central opening of distal shield.

A Gephyrocapsa species with an elevated bridge of two high, paddle-like plates has been
reported (Nishida, 1979; Heimdal & Gaarder, 1981; Kteijne, 1993). It is not clear at this
stage if this is a malformation, a distinct morphotype or a different species.

284
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

Figure 6.4. Papposphaera species, a, P. arctica (HET), TEM

(from Thomsen, 1981). b, P.
arctica (HOL); inset showing detail of a coccolith, TEM
(from Thomsen et ai, 1988). c, P.
obpyramidalis (HET), SEM. d, P. simplicissima (HET), TEM (from Thomsen et ah, 1!
Scale bars: a-d = 2 urn; b (inset) = 200 nm.

285
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

Figure 6.5. Papposphaera and Pappomonas species, a, Papposphaera borealis (HET), SEM.
b, Papposphaera borealis (HOL), TEM (from Thomsen et al., 1991b). c, Papposphaera
borealis (HOL), detail of coccolith, TEM (from Thomsen et al., 1991b). d, Papposphaera
polybotrys (HOL), TEM (from Thomsen, 1980). e, Pappomonas weddellensis, TEM (from
Thomsen et al., 1988). Scale bars: a, b, d = 2 urn; c, e = urn.
1

286
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

Family Papposphaeraceae
Diploid phase motile; coccospheres covered in pappoliths. A central process is often
prominent. Each pappolith is constructed of a rim of two lath types, one small and confined to
the proximal part of the wall, the other vertically expanded; central area with variable
structures. Haploid phase motile, with holococcoliths typically with tower-like forms.

Pappomonas
Coccospheres with up to three types of coccolith: circum-flagellar coccoliths (CFCs); body
coccoliths (BCs); and antapical coccoliths (AACs). Outer rim of coccoliths with angular tops
forming a serrated margin.

Pappomonas weddellensis H.A.Thomsen, in Thomsen et al. (1988: 429, figs 32-34)

Fig. 6.5e

description: Coccospheres 5-6 urn; central area with an axial cross; BCs 1-1.5 um long;
CFC process c. 2-2.5 um high, with a short stem supporting 2 large fan-like blades, one
smaller than the other. AACs much-reduced symmetrical central processes, without a shaft
and smaller distal appendages, sometimes with base-plate calcification of 8 radial bars
instead of 4.

Papposphaera
Heterococcospheres monomorphic or variable; coccoliths elliptical; outer rim elements with
angular tops forming a serrated margin. Species separated mainly on central area and process
structures.

Papposphaera arctica (Manton, J.Sutherl. & Oates) H.A.Thomsen et al. (1991b: 640-641,
figs 5-7, 10) HET
BASIONYM: Turrisphaera arctica Manton, J.Sutherl. & Oates (Manton et al., 1976c: 188-189, pi. 4, figs 24-34)
SYNONYM (original name of HET phase): Papposphaera sarion H.A.Thomsen (1981)

Fig. 6.4a

description: Coccospheres spherical, c. 7 um; coccoliths 1.5-2 um long; central area with an
axial cross; process 2-4 um high, ending distally in 4 subparallel crystallites, each c. 1 um
long; calyx reduced to 4 subparallel rods.

Papposphaera borealis (Manton, J.Sutherl. & Oates) H.A.Thomsen et al. (1991b: 640,
figs 2-4, 9, 12) HET
BASIONYM: Turrisphaera borealis Manton, J.Sutherl. & Oates (Manton et al., 1976c: 187-188, pi. 1, figs 3-
11; pi. 2, figs 12-17; pi. 3, figs 18-23)
SYNONYM (original name of HET phase): Papposphaera sagittifera Manton, J.Sutherl. & McCully (Manton et
al, 1976b)

Fig. 6.5a

description: Coccospheres 4-8 um; coccoliths 1.5-2 um; central areas with an axial cross
and 2-4 additional longitudinal bars; process 2-4 um high, with a small calyx of 4 radially
diverging blades.

Papposphaera obpyramidalis H.A.Thomsen, in Thomsen et al. (1988: 431, figs 35-40)

Fig. 6.4c

description: Coccospheres spherical, c. 5 um; coccoliths c. 1.5 um long; central area with
an axial cross; process 1.5-2.5 um high; low stem supporting a broad cone of 4 large

287
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

elements with rounded distal edges, forming an almost continuous outer layer to the
coccosphere.

Papposphaera polybotrys (H.A.Thomsen) H.A.Thomsen et al. (1991b: 641)

BASIONYM: Turrisphaera polybotrys H.A.Thomsen (1980a: 531, figs 3-5)

The heterococcolith phase has not been observed, but given the large number of
Papposphaera-Turrisphaera combinations that are known, this name was recommended for
both the HET and HOL phases by Thomsen et al. (1991b).

Papposphaera simplicissima H.A.Thomsen, in Thomsen et al. (1988: 443, figs 41-43)

Fig. 6.4d

DESCRIPTION: Coccospheres spherical, c. 4 urn; coccoliths c. 0.5 urn long; central area open;
process absent; rim elements separated so that the wall is discontinuous; crystallites of two
types occur alternately along the scale rim.

Family Syracosphaeraceae
Diploid phase motile; coccosphere completely or largely covered in caneoliths; often
showing dithecatism, with endothecal layer of caneoliths normally conservative in form,
exothecal layer variable, delicate and rarely preserved; and/or modified polar coccoliths,
often forming appendages. Haploid phase motile, with holococcoliths, typically dome-shaped
forms.

Calciopappus
CFCs modified into elongated spines. See Young et al. (2003) for descriptions of species.
Calciopappus sp. was recorded from 57°31'S, south of Tasmania, by Nishida (1986).

Syracosphaera Lohmann (1902)

SYNONYMS: Deutschlandia Lohmann (1912)
Caneosphaera Gaarder & Heimdal (1977)
Gaarderia Kleijne (1993)

Usually dithecate; exothecal coccoliths (XCs) highly variable; endothecal coccoliths are
muroliths with 1, 2 or 3 flanges; the form with 2 flanges often placolith-like; often
differentiated into BCs, CFCs and AACs.

Syracosphaera molischii J.Schiller (1925a: 21, fig. Ka, b)

SYNONYMS: Syracosphaera corrugis Okada & Mclntyre (1977: 21-22, pi. 8, figs 3, 6)

Syracosphaera elatensis Winter, in Winter et al. (1979: 207-208, pi. 13)

Caneosphaera mollischii (J.Schiller) Gaarder, in Heimdal & Gaarder (1981: 44-46, pi. 3)

See also Heimdal & Gaarder (1980) and Kleijne (1993).

Fig. 6.6d

description: BCs 2-4 urn; distal shield flange broad, variably ornamented, usually heavily
ribbed, sometimes with teeth protruding into the central area. Central area an irregular ridge
and/or ring formed by upward growth of laths. CFCs with a prominent spine, smaller and
less calcified than BCs, with narrower flanges. A single AAC usually present, similar to the
BCs but with a stubby spine. XCs large; flange ribbed on distal side, expanding clockwise
into wing (directed apically) c. twice as wide as the rest of the flange.

Several morphotypes can be recognised by their ornamentation.

288
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

Syracosphaera pulchra Lohmann (1902: 134, pi. 4, figs 33, 36a, b, 37) HET
Fig. 6.6a

description: Coccospheres 15-25 pm. XCs dome-shaped; rim subhorizontal, flange-like;

numerous elements with wide slits; central part formed of several cycles of laths. Central
area with a conical depression. BCs 4.5-8 um; mid-wall flange prominent; inner wall-cycle
well developed; central area with 3 concentric cycles of thin radial laths, occasionally with a
small central spine. CFCs similar, with a robust bifurcate spine.

HOL phase synonyms: Calyptrosphaera oblonga Lohmann (1902)

Calyptrosphaera {Daktyletha) pirus Kamptner (1937)

Both holococcolithophore species have been found in combination form with S. pulchra.

Family Umbellosphaeraceae

Diploid phase non-motile; coccospheres with trumpet-like coccoliths. Haploid phase

unknown.

Figure 6.6. Syracosphaera species, a, S. pulchra (HET), SEM. b, S. pulchra (HOL) oblonga
type, SEM (courtesy of M.Knappertsbusch, EMIDAS database), c, S. pulchra (HOL) pirus type,
SEM (courtesy of M.Knappertsbusch, EMIDAS database), d, S. molischii (HET), SEM. Scale
bars: a-d = 2 urn.

289
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

Umbellosphaera
Coccospheres subspherical; coccoliths placolith-like, with a thin distal shield that is greatly
extended. This genus often dominates oligotrophic coccosphere communities.

Umbellosphaera irregularis Paasche, in Markali & Paasche (1955: 97, pi. 3-6)

Fig. 6.7a

DESCRIPTION: Coccoliths concave distally, giving the coccosphere an irregular profile; distal
surface smooth.

Figure 6.7. Umbellosphaera and Wigwamma species, a. U. irregularis, SEM (courtesy of

J.Young. EMIDAS database), b. W. antarctica, TEM. c, W. arctica, TEM. d, W. triradiata,
SEM. Scale bars: a-c = 2 urn; d = urn.
1

290
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

fc«. -Ki

Figure 6.8. Wigwamma and Trigonaspis species, a, W. annulifera (HET), TEM; inset shows
body coccoliths, TEM(from Thomsen et al, 1988). b, W. annulifera (HOL), TEM (from
Thomsen et al., 1988). c-e, T. melvillea. c, Complete cell (type specimen), TEM (from Thomsen
et al, 1988). d, Distal part of coccolith, TEM (from Thomsen et al., 1988). e, Flagellar pole
coccoliths, TEM (from Thomsen et al., 1988). Scale bars: a-c = 1 pm; d, e = 200 nm.

291
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

Genera Incertae Sedis

Wigwamma
Coccoliths hoop-shaped, with a narrow rim and a 'wigwam' of 2-4 rods; rim of 2 concentric
cycles of elements; rim similar in size to Papposphaera.

Wigwamma annulifera Manton, J.Sutherl. & Oates (Manton et al, \9U: 150-152, pi. 4,
fig. 13; pi. 5, figs 14, 15; pi. 6, figs16-21; pi. 7, fig. 21) HET
Fig. 6.8a

description: Coccospheres c. 5 um, dimorphic; coccoliths c. 1-1.5 urn. BCs simple rings;
CFCs with wigwam of 2-4 rods, each with a flange.

HOL phase synonym: Calciarcus cf. alaskensis Manton, J.Sutherl. & Oates (Manton et al,
1977)

Wigwamma antarctica H.A.Thomsen, in Thomsen et al. (1988: 423, figs 9-15)

Fig. 6.7b

description: Coccospheres c. 5 urn, dimorphic; 1 or 2 coccoliths at posterior end with

superstructures 0.8-1.4 um; elsewhere coccoliths 1-1.4 urn. BCs a simple ring; CFCs with a
wigwam of 2-4 rods (2 broad and wing-like, 2 narrow) without flanges; one rod projects as a
spine beyond apex of wigwam; 1 or 2 coccoliths with smaller wigwams can occur at the
posterior end.

Wigwamma arctica Manton, J.Sutherl. & Oates (Manton et al., 1977: 151, pi. 1, figs 1-5;
pi. 2, figs 6, 7; pi. 3, figs 10-12)

Fig. 6.7c

description: Coccospheres 4-6 urn, uniform; coccoliths 1.5-2 um, with a wigwam of 4 rods.

In Arctic regions, each wigwam rod is attached to an enlarged crystallite in the rim; in the
Weddell Sea the rod is attached to a rim crystallite that does not differ from others.

Wigwamma triradiata H.A.Thomsen, in Thomsen et al. (1988: 425, figs 16-20)

Fig. 6.7d

description: Coccosphere 4-6 um, dimorphic. BCs with a low wigwam of 3 rods; CFCs
with a high wigwam of 2 rods, each with a wing.

Holococcolithophores

Cells covered in holococcoliths, each composed of equant microcrystals arranged in planar

arrays. By contrast, heterococcoliths have complex crystals arranged in radial arrays.
Holococcolithophores were formerly placed in the family Calyptrosphaeraceae, but it is now
known that they are formed in the haploid phase of heterococcolith-bearing species.
Nonetheless, for identification purposes, it is convenient to consider holococcolithophores
separately from heterococcolithophores.

Calcidiscus leptoporus (G.Murray & V.H.Blackman) Loebl. & Tappan subsp. ieptoporus
HOL
SYNONYM: Crystallolithus rigidus Gaarder

Fig. 6. Id

292
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

description: Coccoliths with a double layer of crystallites in a hexagonal meshwork array,

upper layer often incomplete; rim 3 crystallites high.

Calcidiscus leptoporus subsp. quadriperforatus (Kamptner) Geisen et al. (2002) HOL

BASIONYM: Syracosphaera quadriperforata Kamptner
SYNONYMS: Syracosphaera (Syracolithus) quadriperforata Kamptner
Syracolithus quadriperforatus (Kamptner) Gaarder

Fig. 6.1b

description: Internal walls of coccoliths define 4-6 pores. Double-layered tube 10-15
where it meets the internal walls; base open. Coccoliths
crystallites high, castellated, higher
2-2.5 urn.

Helicosphaera carteri (G.C.Wall.) Kamptner (1954) HOL

HOL phase synonyms: Syracolithus catilliferus Kamptner [solid variant form]
Syracolithus confusus Kleijne [perforated variant form]

Fig. 6.2c, d

description: Two coccolith variants have been recognised; both occur on single
coccospheres (Cros et al., 2000; Geisen et al., 2000).

Solid variant coccoliths with a flat top showing layered-rhombic fabric; pyramidal boss or
low spine near the centre; rim 1 crystallite wide, showing hexagonal fabric; 5-6 crystallites
high; central opening on proximal side small or absent.

Perforated variant coccoliths with 7-10 circular depressions around the central boss.

Papposphaera arctica (Manton, J.Sutherl. & Oates) H.A.Thomsen et al. (1991b: 640,
figs 5-8) HOL
BASIONYM: Turrisphaera arctica Manton, J.Sutherl. & Oates (Manton et al., 1976c)

Fig. 6.4b

description: Coccospheres spherical, c. 7 um, uniform. Coccoliths c. 1-1.5 um, apple-core-

shaped.

Papposphaera borealis (Manton, J.Sutherl. & Oates) H.A.Thomsen et al. (1991b: 641,
figs 2-4, 9, 11) HOL
BASIONYM: Turrisphaera borealis Manton, J.Sutherl. & Oates (Manton et al., 1976c)

Fig. 6.5b, c

description: Coccospheres elongate, 1.6-8 um, varimorphic. Coccoliths higher at flagellar

pole and not widened distally.

Papposphaera polybotrys H.A.Thomsen (1980a) HOL

BASIONYM: Turrisphaera polybotrys H.A.Thomsen (1980a)

Fig. 6.5d

description: Coccospheres c. 5 urn, spherical, dimorphic. Coccoliths larger at flagellar end,

goblet-shaped; base subcircular; stem narrow, flaring. BCs c. 1 um high. CFCs c. 3 um high,
unilaterally enlarged and closed distally.

Syracosphaera pulchra Lohmann (1902) HOL pirus type

HOL phase synonym: Calyptrosphaera pirus Kamptner; usually ascribed to Daktyletha (Norris, 1985)

Fig. 6.6c

293
 HAPTOPHYTES: ORDER COCCOLITHOPHORALES

DESCRIPTION: BCs tube c. 10 crystallites high; wall fabric tiered; flange basal; distal cover
recessed into a tube, domed with a ring of 8-10 pores around contact with tube; pores
sometimes surrounded by internal ridges. CFCs more elevated and with a pyramidal central
boss.

Syracosphaera pulchra Lohmann (1902) HOL oblonga type

HOL phase synonym: Calyptrosphaera oblonga Lohmann

Fig. 6.6b

description: Coccoliths 2-2.5 um. BCs tube initially flaring, then curving into a convex
cover; proximal surface with a basal flange and 2 concentric rings of crystallites; central
opening large; hexagonal mesh fabric to tube and cover. CFCs more elevated and with a
pyramidal central boss.

Trigonaspis

Coccoliths formed of triangular crystallite groups, c. 0.15 um across, each composed of 3

(?hexagonal) subunits, dimorphic. CFCs tower-like; BCs simple discs.

Trigonaspis melvillea H.A.Thomsen, in Thomsen et al. (1988: 425, figs 23-29)

Fig. 6.8c-e

description: Coccospheres spherical, 3-4 um. CFCs with a slender tube ending distally in a
single distally directed triangular plate. BCs only slightly raised above base-plate.

Wigwamma annulifera Manton, J.Sutherl. & Oates (Manton et al, 1977: 155-156, pi. 8,

figs 22-26) HOL

SYNONYM: Calciarcus cf. alaskensis Manton, J.Sutherl. & Oates (see Thomsen et al., 1991b: 639, fig. 18)

Fig. 6.8b

description: Monomorphic rhombohedral crystallites forming 3 or 4 radial converging

struts attached to a scale with a calcareous rim. Scales c. 0.5 um wide.

294
 7. CHRYSOPHYTES*
Harvey J. Marchant and Fiona J. Scott

The chrysophytes, or golden-brown flagellates, include mostly unicellular, motile and non-
motile protists. Freshwater species are more diverse than marine ones, and many form
characteristic resting cysts during part of their life cycle. These resting cysts (also called
statospores or stomatocysts) have heavily silicified walls, and are thus reasonably resistant to
the dissolution processes of aquatic environments. A number of unidentified cysts found in
Antarctic seawater samples may represent chrysophyte species.

Cells can be naked, covered in silicified or cellulose scales, or bear a chitinous lorica. They are
often spherical to pyriform, but can become distorted following preservation (e.g. in Lugol's
solution). Mixotrophy, when plastid-bearing cells ingest dissolved organic compounds and/or
particulate food, is common among Being nutritionally opportunistic, this
the chrysophytes.
makes them an interesting component of Antarctic microbial communities, particularly during
the winter period. Aspects of chrysophyte taxonomy and biology were documented by Pienaar
(1980) and Kristiansen & Anderson (1996).

Order Ochromonadales

Family Paraphysomonadaceae

Paraphysomonas antarctica E.Takah. (Takahashi, 1987: 158, figs 1 (type), 2-5)

ADDITIONAL REFERENCE: Vors (1992: 64, fig. 3 Id, e)

Fig. 7.1a-c

description: Cells spherical, colourless, 2-4.3 \xm diameter, biflagellate. Flagella unequal;
long flagellum 12.5-27 urn long, with mastigonemes; short flagellum 1.8-4.5 \xm long. Cell
body covered with 1 type of body scale consisting of a basal plate with a central spine; scales
circular, 0.9-1.75 p.m diameter, without a thickened rim or upturned margin; spine
comprising a shaft 0.7-2.55 um long and a thinner apical rod 0.3-0.7 nm long. Cysts
ellipsoidal, c. 5 * 5.6 um (Takahashi, 1987; figs 4, 5).

distribution: Southern Ocean, south of Australia; fast-ice and seawater around East Ongul I.,

East Antarctica (Takahashi, 1987); New Zealand (Moestrup, 1979).

Paraphysomonas antarctica is morphologically similar to P. imperforata (q.v.).

Paraphysomonas butcheri Pennick & K.J.Clarke (1972: 45, figs 1-13)

ADDITIONAL REFERENCES: Takahashi (1987: 163, fig. 12), Vors (1992: 65, fig. 30e), Ikavalko &
Thomsen (1996: 152, figs 3, 4)
Fig. 7.2a-e

description: Cells spherical, 2.4-3.2 um diameter, biflagellate. Flagella unequal; long

flagellum 6-7.5 p.m long, covered with mastigonemes; short flagellum 2.4-3.0 urn long,
smooth. Cell body covered with plate scales and crown scales. Plate scales elliptical, 0.7-0.82
urn long, 0.55-0.63 um wide, mesh-like, with fine perforations, without a central spine or
thickened rim. Crown scales circular, with a fenestrated crown-like superstructure supported
by 4 struts on a simple basal ring.

Class Chrysophyceae.

295
 CHRYSOPHYTES

distribution: Southern Ocean, south of Australia; fast-ice and seawater around East Ongul I.,
East Antarctica (Takahashi, 1987); New Zealand (Moestrup, 1979); a widespread species
(Preisig & Hibberd, 1982a, b); Baltic sea-ice (Ikavalko & Thomsen, 1996).

Paraphysomonas cribosa I. A.N. Lucas (1968: 437, figs 1-5)

Fig. 7.3e

description: Cells ovoid, 3.2-3.8 urn diameter, biflagellate. Flagella unequal, c. 15 um long
and c. 4 um long. Cell body covered with 3 types of scales: plate scales, crown scales and
pyramid scales (isolated scales observed). Plate scales oval, 0.75-0.9 um long, 0.54-0.62 um
wide, built of radial rods supporting an inner oval ring (Fig. 7.3e). Crown scales similar but
bearing an upright crown-like structure supported on 3 or 4 struts to 0.2 um high from an
inner oval ring. Pyramid scales 0.73 um basal diameter, with a central pyramid-like structure
comprising 3 or 4 struts; overall length of scale 1.2-1.62 um.

distribution: Southern Ocean, south of Australia; Plymouth Sound, Great Britain (Lucas,
1968).

Paraphysomonas cylicophora B.Leadb. (Leadbeater, 1972c: 179, figs 1-13)

ADDITIONAL REFERENCE: Vors (1992: 65, fig. 30a, b)

Fig. 7.1d-f

description: Cells spherical, colourless, c. 5 um diameter, biflagellate. Flagella unequal, c.

15 um long and c. 2 um long (Leadbeater, 1972c). Cell body covered with 1 type of scale
comprising a basal plate with a central funnel-shaped superstructure; basal plate circular,
0.6 (am diameter, without a thickened rim; funnel-shaped structure 0.6 um high and c. 0.5 um
diameter at the open end.

distribution: Southern Ocean, south of Australia; type locality, NW of Bergen, Norway

(Leadbeater, 1972c); Baltic Sea (Vors, 1992).

Paraphysomonas imperforata I. A. N.Lucas (1967: 330, pi. la, b, d)

additional references: Preisig & Hibberd (1982: 403, fig. 2A-F), Vors (1992: 64, figs 26e, f,

30f, g)

Fig. 7.3a-c

description: Cells spherical, 3.6-5.1 um diameter, biflagellate. Flagella unequal; long

flagellum 13.5-18 um long, covered with 1.5 um long mastigonemes; short flagellum c. 4.5 um
long. Cells with 1 type of body scale; scales circular, 0.8 um diameter, with a central spine
1-1.3 um long, without a thickened rim.

distribution: Southern Ocean, south of Australia; sea-ice near Casey Station, East
Antarctica (McConville &
Wetherbee, 1983); East Australian waters (Hallegraeff, 1983);
New Zealand (Moestrup, 1979); British waters (Lucas, 1967); Denmark (Thomsen, 1975).

Paraphysomonas imperforata is also known from many freshwater habitats, e.g. lakes in

Greece (Kristiansen, 1980) and Denmark (Wujek & van der Veer, 1976) and a river in Chile
(Durrschmidt, 1980).

Paraphysomonas oligocycla E.Takah. (Takahashi, 1987: 161, figs 8 (type), 9)

additional reference: Vers (1992: 66, fig. 31a-c, e)

Fig. 7.3f

description: Cells spherical to ovoid, 6-6.5 um diameter, biflagellate. Flagella unequal.

Cellbody covered with 1 type of scale, consisting of a basal plate with a central spine; basal
plate

296
 CHRYSOPHYTES

Figure 7.1.Paraphysomonas species, a-c, P. antarctica. a, TEM Whole cell with flagella.
b, TEM Body scale, c, TEM Body scale, d-f, P. cylicophora, SEM Details of body scales.
Scale bars: a = 2 |im; b-f = 0.5 fim.

297
 CHRYSOPHYTES

.
*»

i /

Figure 7.2. Paraphysomonas butcheri. a, SEM Whole cell with flagella. b. TEM Whole cell
with flagella. c, TEM Whole cell with flagella. d, TEM Crown and plate scales, e, TEM Plate

scales. Scale bars: a-c = 2 urn; d, e = 0.5 |im.

298
 CHRYSOPHYTES

Figure Paraphysomonas species,

7.3. a-c, P. imperforata. a, TEM Whole cell with flagella.
b, TEM Whole cell with flagella. c, TEM Scale, d, Paraphysomonas type A, TEM scale.
e, P. cribosa, TEM Plate scale and pyramid scale, f, P. oligocycla, TEM Isolated plate scale.
Scale bars: a, b = 2 urn; c-f = 0.5 fim.

299
 CHRYSOPHYTES

V
< (

rt
Figure 7.4. Paraphysomonas vestita. a, TEM Whole cell with flagella. b, SEM Whole cell
with flagella. c, SEM Detail of body scales, d. TEM Detail of body scales. Scale bars: a, b =
5 jam; c = 2 jam; d = 0.5 \im.

300
 CHRYSOPHYTES

circular, 0.47-0.8 um diameter, ornamentated with 2-4 concentric rings of perforations, with
or without a slightly thickened rim; spine slightly tapering, 0.5-0.9 um long, with or without
a swollen basal part.

distribution: Type locality, fast-ice around East Ongul I., East Antarctica (Takahashi,
1987); Gulf of Finland (Vors, 1992); Arctic Canada (Vors, 1993a).

Details of body scale ornamentation and a comparison with the closely allied Northern
Hemisphere species P. foraminifera I. A. N.Lucas and P. circumforaminifera Wujek were
provided by Takahashi (1987).

Paraphysomonas vestita (A.Stokes) De Saedleer (1929: 177)

BASIONYM: Physomonas vestita A. Stokes (1885: 313)

ADDITIONAL REFERENCES: Takahashi (1987: 163, figs 10, 11), Vors (1992: 66, fig. 30c)

Fig. 7.4a-d

DESCRIPTION: Cells spherical, 5.9-7.5 um diameter, biflagellate. Flagella unequal; long

flagellum c. 18 um long, with conspicuous mastigonemes; short flagellum c. 2 um long. Cell
body covered with 1 type of body scale consisting of a basal plate with a central spine; basal
plate circular, 0.7-0.76 um diameter, with a variably thickened rim; spine 0.1-4.2 um long,
blunt (Fig. 7.4d).

DISTRIBUTION: Coastal waters off Davis Station, East Antarctica; sea-ice around East Ongul I.,

East Antarctica (Takahashi, 1987); Baltic Sea (Vors, 1992).

Most previous records of this widely distributed species come from freshwater habitats
(Takahashi, 1987), with a few from marine environments (Preisig & Hibberd, 1982). The
body scale spines of Antarctic material are somewhat shorter than those of specimens from
other localities (see Takahashi, 1987).

Paraphysomonas type A
Fig. 7.3d

DESCRIPTION: [Only one isolated plate scale was observed.] Scale circular, 0.47 um diameter,
with a marginal ring of 14 perforations and a blunt central spine 0.52 um long.

distribution: Southern Ocean, south of Australia.

The scale seen by us is similar to the freshwater P. circumforaminifera Wujek, a species with
scales 0.4-0.55 jam diameter, patterned with a single marginal ring of 18-22 perforations,
and with a blunt central spine 0.7-0.9 um long. Our material is also reminiscent of
which has 2-4 concentric rings of perforations.
P. oligocycla

Order Parmales

This order comprises two families, the Triparmaceae (with cell walls composed of 8 siliceous
plates) and the Pentalaminaceae (with cell walls composed of 5 siliceous plates) (Booth &
Marchant, 1987, 1988). Cells are difficult to observe by light microscopy because of their
small size. First recognised by SEM, they were tentatively identified as resting stages of
loricate choanoflagellates. Transmission electron microscopy of sectioned organisms showed
them to have a large, bilobed chloroplast, very little storage material and chrysophyte-like
organelles (Marchant & McEldowney, 1986).

Parmales have been reported mainly, but not exclusively, from Antarctic and sub-Arctic
latitudes (Booth & Marchant, 1987) and, more recently, from subtropical waters (Kosman et
ai, 1993). In the Antarctic they are known from the Weddell Sea (Silver et al., 1980; Buck
& Garrison, 1983), Prydz Bay (Marchant & McEldowney, 1986), Lutzow-Holm Bay
(Takahashi et ai, 1986) and Kita-no-seto Strait, north of the Japanese Syowa Station in East

301
 CHRYSOPHYTES

Antarctica (Nishida, 1986). Parmales occur in greatest numbers between 75 and 150 m depth,
atconcentrations in the order of 10" cells per litre (Nishida, 1986; Booth & Marchant, 1987).
They are abundant in siliceous sediments in Prydz Bay (Franklin & Marchant, 1995) and the
Weddell Sea (Zielinski & Gersonde, 1997).

Cell division has not yet been observed in the Parmales, and they may represent stages in an
unknown life cycle (Booth &
Marchant, 1987; Kosman et al., 1993). A diagnostic key to
genera and species was provided by Kosman et al. (1993).

Family Triparmaceae

Tetraparma pelagica B.C. Booth & H.J. Marchant (1987: 248, figs 2 (holotype), 3)

ADDITIONAL REFERENCES: Booth et al. (1981: 76, figs 60-62, as "Cyst IX"), Marchant &
McEldowney (1986: 54, fig. 5), Kosman et al. (1993: 123, fig. 31)
Fig. 7.5a, b

DESCRIPTION: Cells planktonic, solitary, non-motile, spherical, 2.2-2.8 urn diameter. Cell
wall comprising 8 siliceous plates: 4 shield plates and 4 triradiate plates,
all fitting edge to

edge, with raised rims; all plates with branching veins, with or without surface papillae.
Shield plates circular, 1.7-2.2 um diameter, with a central projection to 3.5 urn long.
Triradiate plates with arms c. 1.3 um long. Papillae arranged in rows radiating from the
centre, 10 per micrometre, variously developed on different specimens.

distribution: Surface waters of Prydz Bay, East Antarctica; North Pacific, Bering Sea,
eastern equatorial Pacific, Greenland and Finland (Kosman et al., 1993).

Triparma columacea B.C. Booth subsp. alata H.J. Marchant, in Booth & Marchant
(1987: 251, figs 15 (holotype), 16-18)

ADDITIONAL REFERENCES: Booth et al. (1981: 68, figs 31-38, as "Cyst IV"), Kosman et al. (1993:
124, figs 38-43)

Fig. 7.5c, d

description: Cells planktonic, solitary, non-motile, subspherical or hemispherical, 3.5-4.2 um

diameter. Cell wall comprising 8 keeled siliceous plates: 3 shield plates, 1 ventral plate.
1 triradiate plate and 3 oblong girdle plates; junctions between plates distinct; all plates with

coarse venation, lacking papillae. Shield plates 1.6-2.2 um diameter, flat to slightly convex,
radially veined, with (1-) 2 (-4) central knobs. Ventral plate 2.8-3.5 um diameter, conical or
hemispherical, radially veined, with a smooth slightly raised margin; central process with 1^4
lobes. Girdle plates with a wing-like undulating keel, with a narrow raised margin and
perpendicular veins on either side of keel. Triradiate plate with a triradiate keel, most
pronounced midway along the arms.

DISTRIBUTION: Prydz Bay, East Antarctica; sub-Arctic Pacific (Booth & Marchant, 1987).

Triparma laevis B.C. Booth subsp. pinnatilobata H.J. Marchant, in Booth & Marchant
(1987: 255, figs 34 (holotype), 35, 36)

Fig. 7.6a

description: Cells planktonic, solitary, non-motile, spherical, 3-3.6 um diameter. Cell wall
comprising 8 siliceous plates: 3 shield plates, ventral plate, 1 keeled triradiate plate keel and
1

3 girdle plates; junctions between plates distinctively raised; all plates lacking veins and
papillae. Shield plates circular, 1.7-2 um diameter, slightly convex, smooth, with a raised
margin, 1-3 short processes sometimes present in centre of plate. Ventral plate convex, 2.4-
2.6 um diameter, with a slightly raised rim, an irregular spathulate central process and/or a
concentric ring of processes that may be dichotomously branched. Triradiate plate arms 1.4-
1.7 um long; triradiate keel sometimes well developed, with a slightly raised margin; plate

302
 CHRYSOPHYTES

Figure 7.5. Tetraparma and Triparma species, a. b, Tetraparma pelagica, SEM. c. d,

Triparma columacea subsp. alata, SEM. Scale bars: a-d = p.m.
1

303
 CHRYSOPHYTES

Figure 7.6. Triparma and Pentalamina species, a, T. laevis subsp. pinnatilobata, SEM. b, T.
laevis subsp. ramispina, SEM. c, T. strigata, SEM. d, P. corona, SEM. Scale bars: a-d = 1 urn.

304
 CHRYSOPHYTES

slightlyconvex. Girdle plates with pronounced keels; processes irregularly shaped,

sometimes dichotomously branched, c. 2 pm long. Ornamentation of plates, including the
number and placement of processes and their length, can vary.
distribution: Widely distributed in Antarctic waters; Weddell Sea (Silver et al., 1980);
south of Australia (Nishida, 1986); Prydz Bay, East Antarctica, (Marchant & McEldowney,
1986).

Triparma laevis B.C. Booth subsp. ramispina H.J. Marchant, in Booth & Marchant
(1987: 256, figs 37 (holotype), 38)

ADDITIONAL REFERENCES: Silver et al. (1980: 212, fig. lb), Nishida (1986: 66, pi. 1, fig. 6),
Kosman et al. (1993: 124, figs 29, 30)

Fig. 7.6b

description: Cells planktonic, solitary, non-motile, spherical, 2.5-3.2 um diameter. Cell

wall comprising 8 smooth and ornamented siliceous plates: 3 shield plates, 1 ventral plate,
1 triradiate plate and 3 girdle plates; junctions between plates distinctively raised; all plates
smooth. Shield plates slightly convex, with raised margins, 1.8-2 um diameter, with or
without central processes that can be rounded, spathulate or dichotomously branched.
Ventral plate convex, with a narrow raised rim, with or without central irregularly branched
processes 1.8-2.6 um long. Triradiate plate with or without an undulating triradiate keel;
arms of plate 1.7-1.9 pm long; margin of plate raised as a narrow smooth or irregular rim.
Girdle plates with long dichotomously branched spines to 8 um long.

distribution: Prydz Bay, East Antarctica; Terra Nova Bay; other Antarctic waters (Booth &
Marchant, 1987).

Although a clear distinction can be made between subspecies ramispina and pinnatilobata,
these organisms exhibit considerable structural variability. In our experience the two
subspecies are the most abundant Parmales in Antarctic waters.

Triparma strigata B.C. Booth, in Booth & Marchant (1987: 256, figs 40 (holotype), 41,
42)

additional reference: Booth et al. (1981: 68, figs 24-30, as "Cyst III")

Fig. 7.6c

description: Cells planktonic, solitary, non-motile, hemispherical, 3.1-3.8 pm diameter,

c. 2.8 pm long; dorsal surface convex; ventral surface flat. Cell wall comprising 8 siliceous
plates: 3 shield plates, 1 ventral plate, triradiate plate and 3 girdle plates; junctions
1

between plates usually indistinct; all plates bearing similar processes to 1.5 pm long, straight
or forked, round in transverse section. Shield plates round, rarely concave, 1.8-2.8 pm
diameter, with delicate radiating striae. Ventral plate flat, 2.4-3.3 pm diameter, with delicate
radiating striae. Triradiate plate without a keel; arms 1.4-2.3 pm long. Girdle plates with
scattered processes, each with 2 straight or forked spines to 9 pm long.

distribution: Sub-Arctic Pacific and Antarctica (Booth & Marchant, 1987).

Little morphological variation has been observed in this species, but Booth & Marchant
(1987) reported that Antarctic cells had straight spines on the girdle plates in contrast to the
branched spines of sub-Arctic material.

Family Pentalaminaceae

Pentalamina corona H.J. Marchant, in Booth & Marchant (1987: 258, figs 6 (holotype),
7)

ADDITIONAL REFERENCES: Buck & Garrison (1983: 1275, fig. 35, as a siliceous cyst), Takahashi
et al. ( 1 986: 87, figs 17, 18, as "Cyst 4A")

305
 CHRYSOPHYTES

Figure 7.7. Meringosphaera mediterranea. a, SEM Partially collapsed cell, b, TEM Entire
cell, c, d, TEM Details of spines. Scale bars: a, b = 5 urn; c, d = 2 (am.

306
 CHRYSOPHYTES

Fig. 7.6d

description: Cells planktonic, solitary, non-motile, spherical or subspherical, 5-5.8 |im

diameter. Cell wall comprising 5 siliceous plates: 3 round plates and 2 triradiate plates.
Round plates of unequal size, 3.6-4.6 urn diameter, slightly convex, with or without rounded
spathulate or dichotomously branched processes (3 urn long) arising from the rims and centre
of the plate. Triradiate plates with raised margins, some with processes 1.5 urn long, with or
without a central process 1.8 um long. Margins between triradiate and shield plates distinct;
margins between triradiate plates difficult to observe. Ornamentation of the plates can vary.

distribution: Prydz Bay, East Antarctica; Terra Nova Bay, Ross Sea; Weddell Sea (Silver et
al.,1980; Buck & Garrison, 1983); Kita-no-seto Strait, near Syowa Station, East Antarctica
(Takahashiefa/., 1986).

Order Synurales

Family Pleurochloridaceae

Meringosphaera mediterranea Lohmann (1902: 89)

ADDITIONAL REFERENCES: Moestrup (1979: 64, figs 2, 6), Buck & Garrison (1983: 1265, figs 2,
3), Chretiennot-Dinet (1990: 43, figs 140-142)

Fig. 7.7a-d

DESCRIPTION: Cells solitary, non-motile, readily identified by wavy spines projecting from a
spherical body. Cell body to 5-9 um diameter, covered with elliptical plate scales c. 0.4 x
0.8 um, each bearing a central spine. Spine c. 13-15 um long, tapering, with short lateral
barbs directed towards the apex.

DISTRIBUTION: Weddell Sea (Buck & Garrison, 1983); Southern Ocean, south of Australia;
East Australian Current (Hallegraeff, 1983); New Zealand (Moestrup, 1979); Greenland,
Great Britain, Denmark, Adriatic, Gulf of Aqaba, Indian Ocean and sub-Arctic Pacific Ocean
(Booth et al, 1982).

307
 8. PRASINOPHYTES*
Harvey J. Marchant

Originally placed within the Class Chlorophyceae, these small, solitary or colonial cells were
segregated due to having layers of organic scales covering the body and flagella (see Boney,
1970; Norris, 1980). The class, formally typified by Moestrup & Throndsen (1988) based on
the genus Prasinocladus, comprises two orders and some 16 genera. Flagellate forms possess
2 or 4 (exceptionally 6 or 8) anteriorly inserted flagella. Typically, each cell has a single
cup-shaped or lobed chloroplast that is green, yellow, gold or olive, with 1 or more large
pyrenoids. The scales are formed within the Golgi apparatus, later being released to the
surface of the cell. They are distinguishable by SEM or TEM and are finely and distinctively
patterned, this patterning being useful for the recognition of species. The non-motile
vegetative stage of prasinophyte flagellates, the phycoma stage, is often reported.

Prasinophytes are commonly a minor component of the Antarctic phytoplankton with high
spatial and temporal variability in Antarctic waters (Marchant, 1993). Buck & Garrison
4
(1983) recorded c. 0-3 * 10 cells per litre, while Davidson & Marchant (1992a) reported
1.6 x 10 cells per litre. Bird & Karl (1991) documented a massive bloom of a Pyramimonas
5

species in the northern Gerlache Strait with concentrations of chlorophyll a reaching 25 ug

5
per litre and cell numbers exceeding 7 * 10 per litre.

Order Chlorodendrales

Family Halosphaeraceae

Pterosperma parallelum Gaarder (1938: 68, fig. 8a-c)

ADDITIONAL REFERENCE: Throndsen (1997: 664, pi. 12)

Fig. 8.1a

description: Phycoma stage: Cells spherical, 9-20 urn diameter; walls with low curved
parallel alae.

distribution: Southern Ocean, near the Polar Front.

Pterosperma cf. polygonum Ostenf., in Ostenfeld & Schmidt (1902: 141)

ADDITIONAL REFERENCE: Throndsen (1997: 664, pi. 12)

Fig. 8.1b

description: Phycoma stage: Cells spherical, 10 urn diameter (13 urn including alae); walls
with low curved parallel alae in a hexagonal arrangement.

distribution: Coastal waters, Davis Station, East Antarctica.

Material from the Antarctic resembles P. polygonum Ostenf, which has a cell body of 30 jam
diameter and alae in a pentagonal pattern (Throndsen, 1997: 664, pi. 12). Members of the
genus were discussed by Parke et al. (1978).

Class Prasinophyceae.

308
 PRASINOPHYTES

Figure 8.1. Pterosperma and Pyramimonas species, a, Pterosperma parallelum, SEM.

b, Pterosperma cf. polygonum, SEM. c—e, Pyramimonas australis. c, SEM Solitary cell
(from Moro et al., 2002). d, TEM Underlayer scales (from Moro et al., 2002). e, TEM Box
scale, frontal view with typical perforations (from Moro et al., 2002). Scale bars: a, b = 5
Urn; c =1 (im; d = 500 nm; e = 100 nm.

309
 PRASINOPHYTES

Figure 8.2. Pyramimonas gelidicola. a, LM Solitary motile cells, b, LM Dividing cells.

c. SEM Cyst, d, SEM Motile cell; note dislodged scales and distended lipid bodies on left.
e, TEM Motile cell. Scale bars: a, b = 10 fim; c-e = 2 ^m.

310
 PRASINOPHYTES

Pyramimonas
Sym & Pienaar (1993) discussed the diagnostic characteristics of the green flagellate
Pyramimonas. Cells have either 4 or 8 flagella of equal length emerging from the anterior
end of the cell. Up to 7 types of non-mineralised scales cover the cell body and the flagella,
and these scales are diagnostic at the species level.

Pyramimonas australis Andreoli & Moro, in Moro et al. (2002: 105, figs 1-17)

Fig. 8.1c-e

description: [after Moro et al. (2002)] Cells ovoid, to 8-10 urn long, 5 6 um diameter.
Body scales of 3 types: an outer rather loose layer of crown scales, a middle layer of closely
fitting box scales and an inner layer of smaller scales. Crown scales c. 180 nm across the
base, c. 200 nm high, with a central spine and 4 arching ribs attached near the spine tip and
extending down to the base-plate. Box scales square, c. 270 nm on each side, with
16 perforations forming indistinct square subunits. Inner scales c. 150 nm long. Flagella thin,
c. 8.5 um long, with an outer layer of limulus scales and underlying rows of pentagonal

scales; limulus scales c. 250 nm long, with irregular perforations; pentagonal scales c. 40 nm
diameter, with a central boss. Cells with a single lobed chloroplast surrounding a central
pyrenoid. A solitary posteriolateral eyespot is located near the pyrenoid.

Cysts spherical, 9-10 um diameter, with a uniform covering of a single scale type not seen
on the vegetative (flagellate) cell; scales spine-like, with a distinctive stellate tip.

distribution: Type locality, in sea-ice, Terra Nova Bay, Ross Sea (Moro et al, 2002).

Pyramimonas gelidicola McFadden, Moestrup & Wetherbee (1982: 107, fig. 19)

ADDITIONAL REFERENCES: van den Hoff et al. (1989: 448, figs 2-19); Davidson & Marchant
(1992b), McMinn & Hodgson (1993: 936, fig. 4)

Figs 8.2a-e; 8.3a-c

DESCRIPTION: Cells subcylindrical to conical, to 8-18 um long, 7-9 pm diameter. Body

scales of 3 types: an outer rather loose layer of crown scales, and an inner layer of closely
fitting box scales and smaller footprint scales. Crown scales c. 300 nm across the base,
c. 300 nm high, with 4 centrally connected arches. Box scales square, c. 300 nm on each

side, with perforated walls c. 80 nm high. Footprint scales positioned among the box scales,
c. 80 nm long. Flagella thin, marginally longer than the cell body, with an outer layer of

limulus scales and underlying rows of pentagonal scales; limulus scales 400-500 nm long,
with irregular perforations; pentagonal scales c. 60 nm diameter, with a central boss. Cells
with a single lobed chloroplast, with 1 parietal pyrenoid opposite a pair of eyespots.

Cysts spherical, 1 1-20 pm diameter, with a uniform covering of a single scale type not seen
on the vegetative (flagellate) cell; scale base-plate similar to that of vegetative cell box
scales but with a horn-like elongation on each of the 4 corners.

distribution: Coastal waters and fast-ice of the Vestfold Hills, East Antarctica; Southern
Ocean, south of Australia.

The five scale types of vegetative cells, discernable by TEM and SEM, were described and
illustrated in the type description (McFadden et al, 1986). McMinn & Hodgson (1993)
reported this species as the "common dominant green phytoflagellate after sea-ice break-
to
out in early summer" in a fjord in the Vestfold Hills area, co-dominating the midsummer
phytoplankton bloom with another small flagellate, Geminigera cryophila. Davidson &
Marchant (1992b) also recorded P. gelidicola in peak abundance with a Phaeocystis bloom.

Cyst morphology and ultrastructure have been described in detail by van den Hoff et al.
(1989). They also documented the developmental changes that occur during the production
of cysts from vegetative cells.

311
 PRASINOPHYTES

Pyramimonas tychotreta Daugbjerg (2000: 161, figs 2-48)

ADDITIONAL REFERENCE: Daugbjerg et al. (2000: 199-209)

Fig. 8.3d, e

DESCRIPTION: [after Daugbjerg et al. (2000)] Cells 8-12 um long, 6-8 um diameter, with
slightly convex sides. Body scales of 4 types: an outer rather loose layer of crown scales, a
middle layer of closely fitting box scales, an inner layer of small footprint scales, and an
underlayer of minute square scales around the flagellar pit. Crown scales c. 300 nm across
the base; scale base with 4 ribs in a cross supporting a central strut that connects distally
with 4 curved arms arising from the scale base edges. Box scales square, c. 300 nm on each
side, c. 80 nm high, ornamented with parallel striations and scattered perforations. Footprint
scales 70 nm long, 20 nm wide, dispersed between the box scales. Flagella thin, slightly
longer than the cell body, with an outer layer of limulus scales and an underlying rows of
pentagonal scales and 2 rows of hair-shaped scales; limulus scales c. 313 nm long (including
apical spine); pentagonal scales c. 40 nm diameter; hair scales c. 1.3 um long. Cells with a
single cup-shaped chloroplast lobed anteriorly. Two eyespots in adjacent chloroplast lobes
about halfway down the cell.

Cysts spherical, c. 10 um diameter, with a uniform covering of a single scale type not seen

on the vegetative (flagellate) cell; scales spine-like, c. 60 nm high; scale base-plate similar to
that of vegetative cell box scales, 150 nm wide, but with a horn-like elongation 300-500 nm
high with a terminal hook on each of the 4 corners.

distribution: Type locality, Weddell Sea, 60°S 49°W (Daugbjerg, 2000); Ross Sea
(Daugbjerg et al., 2000).

Order Mamiellales

Family Mamiellaceae

Mantoniella antarctica H.J.Marchant, in Marchant et al. (1989: 168, figs 4 (holotype),

6, 12-21,23-25)

Fig. 8.4a, b

DESCRIPTION: Cells spherical to slightly ovoid, 2.8-5 nm diameter, with 2 unequal flagella
arising from anterior part of cell. Cell body and flagella covered with scales of different
types. Body scales virtually flat, of 2 types arranged in separate layers, with smaller scales
overlain by larger ones; inner (smaller) scales like those of the flagella; outer (larger) scales
(280-) 360 (-470) nm diameter, with the same arrangement of concentric threads as smaller
scales but with only 6 radial spokes. Long flagellum 8.6-21.3 um long; short flagellum c. 1
um long. Flagella scales arranged in rows, flat, scalloped in outline, (200-) 240 (-270) nm
diameter, comprising 7 radial spokes and 3 concentric threads; 2 inner concentric threads
forming a pair, outer concentric thread attached to radial spokes close to the junctions of the
spokes producing the scalloped outline; flagella also sparsely decorated with hair-scales 0.51
um long and 30 nm diameter, tapering proximally, with a delicate cross-banded structure
with a periodicity of c. 12 nm.

distribution: Pack-ice, Weddell Sea (Marchant et al., 1989).

The terminal tuft of hair-scales on the longer flagellum of M

squamata (below) is sometimes
found on M. antarctica (Marchant et al, 1989). The two species are difficult to distinguish
from one another without SEM preparations.

Mantoniella squamata (Manton & Parke) Desikachary (1973: 447, pi. 1, fig. 1)

BASIONYM: Micromonas squamata Manton & Parke (1960: 293, figs 1-29)

312
 PRASINOPHYTES

V
3qC V— -•

Figure 8.3. Pyramimonas species, a-c, P. gelidicola. a, SEM Dividing cell, b, TEM Scale
types:box (bx), limulus (li), footprint (ft), pentagonal (pt) and crown (cr). c, TEM Cyst scale
with hooked and knobbed ends, d, e, P. tychotreta. d, TEM Motile cell (from Daugbjerg,
2000). e, TEM Box scales (from Daugbjerg, 2000). Scale bars: a = 2 um; b = 500 nm; c, e =
200 nm; d = 5 um.

313
 PRASINOPHYTES

Figure 8.4.Mantoniella species, a, b, M. antarctica. a, SEM Whole cell (from Marchant et ai,
1989). b, SEM Flagellum and flagellar scales (from Marchant et ai, 1989). c, d, M. squamata.
c, SEM Whole cell (from Marchant et ai, 1989). d, SEM Flagellum with discoid scales and

hair scales (arrow) (from Marchant et ai, 1989). Scale bars: a, c =

1 urn; b, d = 500 nm.

314
 PRASINOPHYTES

ADDITIONAL REFERENCES: Barlow & Cattolico (1980: 321, figs 1-30), Hallegraeff (1983: 499,
fig. 25a, b), Marchant et al. (1989: 168, figs 1-3,7-11,22)

Fig. 8.4c, d

description: Cells 2.7-3.5 [im diameter (dried preparation), with 2 unequal flagella arising
from anterior part of cell. Body scales ( 140—) 240 (-330) nm diameter, with a spider's web
pattern of fibrils and a small central protuberance, with 7 radial spokes, concentric ribs not
regularly spaced, gaps seen around central protuberance. Long flagellum (10-) 22 (-30) urn
long; short flagellum c. 1 urn long. Flagella scales arranged in rows, apparently attached by
their rims, flat, discoid, 0.18-0.26 um diameter, patterned like body scales; flagella also
sparsely decorated with hair-scales that can form a terminal tuft on the long flagellum; hair-
scales c. 0.5 um long, 30 nm diameter, tapering proximally, with a delicate cross-banded
structure with a periodicity of c. 12 nm.

distribution: Pack-ice and land-fast ice, Weddell Sea (Marchant et al, 1989); Great Britain
(Manton & Parke, 1960); eastern Australia (Hallegraeff, 1983); Denmark (Manton &
Leadbeater, 1974); Greenland (Thomsen, 1982).

Cell dimensions and flagellar lengths are variable (Marchant et al, 1989). Scales of
Antarctic specimens are the same size as those reported for the type material from Great
Britain, but they differ structurally. Scales of the latter have 8 radial spokes while the
Antarctic cells have 7. Moreover, concentric ribs are ±regularly spaced in the type material,
less so in Antarctic populations.

315
 9. CHLOROPHYTES
Fiona J. Scott

Unicellular, aquatic chlorophytes usually have the following attributes: cells often rounded,
ovoid or lobed, with or without a cell wall, with 1, 2 or 4 flagella and a bright green colour;
1 chloroplast containing an eyespot and a pyrenoid. Although some species are found in
freshwater, soil and snow habitats in the Antarctic, unicellular chlorophytes are not a
conspicuous component of Antarctic marine ecosystems.

Order Volvocales

Family Chlamydomonaceae

Polytoma papillata Pascher (1927: 577)

ADDITIONAL REFERENCES: Ves (1992: 36, figs 17a-d, 19.1, 2), Iksalko & Thomsen (1997: 234,
pl. 3, fig. 31)

Fig. 9.1a, b

description: Cells ovoid, 18-25 um long, 8-15 um diameter, biflagellate, with a conspicuous
hemispherical papilla at the anterior end. Cell surface finely papillate. Flagella equal,
homodynamic. Cells heterotrophic, the chloroplast being replaced by a colourless leucoplast.
distribution: Coastal waters off Davis Station, East Antarctica; Baltic sea-ice (Vb6, 1992).

Cells found in the present investigation were comparatively small (4.5-5.5 um long), but
otherwise fit well with the description of Polytoma papillata sensu Ves (1992).

Figure 9.1. Polytoma papillata. a, SEM Whole cell, b, TEM Cell. Scale bars: a, b= 5 um.

Class Chlorophyceae.

316
 10. CRYPTOPHYTES*
Fiona J. Scott and John van den Hoff

Cryptophytes are rather delicate flagellates with autotrophic and heterotrophic representatives.
Because of their fragile structure, they are often poorly preserved and, therefore, are
frequently overlooked in plankton studies. Cells are comparatively small, 3-50 urn long, and
they are most abundant in cold or deep waters (Graham & Cox, 2000). Pigment studies have
shown that cryptophytes are widespread in Antarctic waters, but are only rarely abundant (see
Marchant, 1993). Heterotrophic cryptophytes have not yet been recorded from the Antarctic.

Order Cryptomonadales

Family Cryptomonadaceae

Geminigera cryophila (D.L.Taylor & C.C.Lee) D. R.A.Hill (1991a: 181, fig. 2)

BASIONYM: Cryptomonas cryophila D.L.Taylor & C.C.Lee (1971: 270, figs 1 (holotype), 2-18)

ADDITIONAL REFERENCES: Brett & Wetherbee (1986: 23); Archer et al. (1996b: 184)

Fig. 10.1a, b

DESCRIPTION: Cells characteristically ovate and flattened, tapering towards posterior end, 15-
17 urn long, 8-10 um wide, 3-5 um thick, biflagellate; cell surface "warty" or "scalar/
fibrillar" (Hill, 1991a), with a longitudinal furrow; gullet subapical and sack-like. Flagella
equal or subequal, c. 8 um long, covered with fine hairs, arising from the gullet opening.
Cells with a single slightly lobed parietal chloroplast, with 2 conspicuous reniform pyrenoids
on opposite lobes of chloroplast; nucleus situated in the posterior end of the cell.

distribution: Coastal waters near Davis Station, East Antarctica; beneath pack-ice in the
Weddell Sea (Taylor & Lee, 1971).

Hill (1991a) described the distinctive, "warty" or "scalar/fibrillar" appearance of the cell
suface caused by a "prolific accumulation of lipid droplets in the peripheral cytoplasm".

Cryptophyte sp.

Fig. 10.1c, d

description: Cells characteristically tear-shaped, tapering sharply towards the posterior end,
6.5-9 um long, 3.2-5.5 um wide, biflagellate; cell surface of hexagonal plates, each plate
c. 0.7 um across, with raised rims; cells with a longitudinal furrow; gullet subapical and

sack-like, c. half cell body length. Flagella equal or subequal, (5-) 7-10.5 um long, covered
with fine hairs, arising from the gullet opening. Cells with a single parietal chloroplast;
nucleus situated in the posterior end of the cell.

distribution: Coastal waters and hypersaline lakes near Davis Station, East Antarctica.

This organism is likely to be a new cryptophyte genus (D. R.A.Hill, pers. comm.). Pigment
analysis has demonstrated the presence of chlorophyll a, alloxanthin and Cr-phycoerythrin
(Wright, unpublished data).

Class Cryptophyceae.

317
 CRYPTOPHYTES

Figure 10.1. Geminigera cryophila and an indeterminate cryptophyte. a, b. Geminigera

cryophila. a, SEM Dorsal view of cell, b, SEM Ventral view of cell showing gullet and
emergent flagella. c, d, Cryptophyte sp. c, SEM Dorsal view of cell with hexagonal plates of
the periplast, d, SEM Ventral view of cell showing gullet and emergent flagella. Scale bars: a-
d = 5 urn.

318
 ll.EUGLENOIDS*
Fiona J. Scott

Euglenoid cells are solitary, with 2 heterodynamic flagella arising from an anterior canal
(sometimes only one emerges). They have pronounced metaboly (mobility and the ability to
change shape), they possess pellicular striations or bands (often helical) that are usually
visible by LM, and they have an orange-red eyespot outside the chloroplast (in contrast to
chlorophytes) at the base of the canal. Species identification relies heavily on morphology
and the availability of living material, with ultrastructural observations verifying certain
attributes.

There are few reports of euglenoids from Antarctica. Three unidentified species were
recorded from sea-ice near Davis Station, East Antarctica (Archer et o/., 1996b) and two
(including Petalomonas sp.) were found in pack-ice assemblages in the Weddell Sea
(Garrison & Buck, 1989b).

The zoological classification scheme used by Hausmann & Htilsmann (1996) refers to the
Class Euglenophyceae as Phylum Euglenozoa and includes the Subphyla Euglenida and
Kinetoplasta. Here, we have dealt with the latter as Protista Incertae Sedis, Superclass
Kinetoplasta (see p. 460).

Order Sphenomonadales

Family Sphenomonadaceae

Anisonema prosgeobium Skuja (1939: 123, pi. 10, figs 16, 17)

ADDITIONAL REFERENCE: Vors (1992: 41, fig. 18i-k)

Fig. 11.1a, b

description: Cells heterotrophic, usually elongate, slightly compressed, 25-60 um long, 14-
25 um wide, biflagellate. Flagella unequal, heterodynamic, both emerging from an apical canal;
the shorter thicker flagellum directed anteriorly; the longer flagellum looped at the point of
emergence and then directed back along the cell body. Cell surface with a smooth pellicle;
pellicular strips not conspicuous. Cells have a gliding motion with distinctive reversal
behaviour.

distribution: Southern Ocean, south of Australia; Arctic Canada and the Baltic (Vors,
1992).

"Anisonema sp." was recorded from pack-ice assemblages in the Weddell Sea by Garrison &
Buck (1989b).

Order Eutreptiales

Family Eutreptiaceae

Eutreptiella cf. braarudii Throndsen (1969: 271)

ADDITIONAL REFERENCES: Daugbjerg & Vers (1992: 24, fig. 10(5-6)), Walne et at. (1986: 114)

Fig. 11.2a h

Class Euglenophyceae or Subphylum Euglenozoa.

319
 EUGLENOIDS

Figure 11.1. Anisoma and Eutreptiella species, a, b, Anisoma prosgeobium, LM. c-g,
Eutreptiella sp. c, d, LM Solitary cells, e-g, SEM Solitary cells with striated pellicle. Scale
bars: a-g = 10 urn.

320
 EUGLENOIDS

Figure 11.2. Eutreptiella cf. braarudii. a-d, LM Metaboly of cell, e, SEM Elongated cell.
f, SEM Flagellated cell, g, SEM Spherical cell, h, SEM Detail of flagella and pellicle. Scale
bars: a-h = 10 \xm.

32:
 EUGLENOIDS

DESCRIPTION: Cells with pronounced metaboly, 65-95 urn long, c. 16 urn wide at the widest
point, biflagellate.Long flagellum 60 um long; short flagellum 38 urn long. Pellicular bands
15 in 10 um, helical. Reservoir system small.

distribution: Coastal waters off Davis Station, East Antarctica; Norway (Throndsen, 1969).

Eutreptiella sp.

Fig. 11.1 c-g

DESCRIPTION: Cells with pronounced metaboly, 21-32 urn long, 1-20 um wide at the widest
1

point, biflagellate. Long flagellum 36 um long; short flagellum 20 um long. Pellicular bands
9 in 10 um, straight. Reservoir system small.

Figure 11.3. Postgaardi mariagerensis. a. b. LM Solitary cells, c. d, SEM Cells with

emergent flagella and pellicle covered in rod-shaped bacteria. Scale bars: a-d = 10 um.

322
 EUGLENOIDS

distribution: Coastal waters off Davis Station, East Antarctica.

The material collected appears to be a species of Eutreptiella due to its unequal and
anteriorly directed flagella. However, the striations are comparatively coarse (9 in 10 um)
and are straight, not helical as is characteristic of this genus.

Postgaardi mariagerensis Fenchel, C.Bernard, Esteban, BJ.Findlay, P.J.Hansen &

Iversen (1995: 72, figs 15c, 16a-c)

ADDITIONAL REFERENCE: Simpson et al. (1996: 213, figs 2-25)

Fig. 11.3a-d

DESCRIPTION: Cells colourless, ovoid, with somewhat pointed ends, 16-25 um long, 8-10 um
diameter, biflagellate. Flagella equal, c. 2.5 times cell body length, curved or helical,
emerging from a large anterior depression. Cell body covered with longitudinally arranged
rod-shaped bacteria (Fig. 1 1.3c). Living cells swim slowly, spiralling around their long axis.

distribution: Ellis Fjord, near Davis Station, East Antarctica; Mariager Fjord, Denmark
(Fenchel et al, 1995).

Electron microscope studies have demonstrated that P. mariagerensis does not possess a
(Simpson et al, 1996), a feature typical of the euglenoids. It is currently regarded as
pellicle
Euglenozoa incertae sedis until further studies clarify its taxonomic position (Simpson et al,
1996).

323
 12. CYANOPHYTES*
Harvey J. Marchant

Cyanobacteria typically occur in low numbers in Antarctic waters (Marchant et al., 1987a).
Chroococcoid cyanobacteria have been found in sea-ice communities (Waterbury et al.,
1979; Walker & Marchant, 1989; Mathot et al, 1991; Gradinger & Lenz, 1995), and have
been reported in concentrations of up to 140 cells per ml in the water column at an Antarctic
coastal site (Walker & Marchant, 1989). Thus, they represent a minor source of carbon for
higher trophic levels in Antarctic waters.

Order Chroococcales

Synechococcus sp.

ADDITIONAL REFERENCES: Marchant et al. (1987a: figs 1, 3), Walker & Marchant (1989: 193)

Fig. 12.1a, b

description: Cells spherical, 0.8-1.4 urn diameter, solitary or in loose clumps 5-20 um
diameter, with bright yellow-orange autofluorescence. Outer layer of cell wall sometimes
patterned. Cells dividing by binary fission.

Figure 12.1. Synechococcus sp. a, LM Solitary and dividing cells appear yellow-green under
blue epifluorescence microscopy, b, TEM Cell dividing by binary fission (from Marchant et
al., 1987). Scale bars: a = 2 urn; b = 100 nm.

Class Cyanophyceae. Also known as cyanobacteria and blue-green algae.

324
 CYANOPHYTES

distribution: Coastal waters of the Vestfold Hills, East Antarctica (Walker & Marchant,
1989).

The differences in patterning of the cell wall of Antarctic material, observed by TEM, may
represent different strains of Synechococcus (Marchant et al, 1987a). While occurring at
concentrations of about 1-10 million cells per litre in temperate and tropical waters, the
concentration of cyanobacteria in Antarctic waters, where the temperature is less than 2°C, is
only in the hundreds or thousands of cells per litre (Marchant et al., 1987a; Walker &
Marchant, 1989).

325
 13. CHOANOFLAGELLATES*
Harvey J. Mar chant

The choanoflagellates are a well-defined group of unicellular, marine and brackish water
The protoplast is usually colourless and bears a single anteriorly directed flagellum
protists.
surrounded by a ring or collar of tentacles used to trap food particles. Three families are
distinguished on the basis of differences in periplast structure. In the family Codonosigidae,
the protoplast can be either naked or have a fine, often indistinct sheath. Members of the
Salpingoecidae (not known from Antarctic waters) possess a firm theca that is visible by LM
or EM. Species of the family Acanthoecidae have a protoplast that is surrounded by a lorica,
or basket, constructed of costal strips (siliceous cylindrical or flattened rods). The costal
strips can be longitudinally, transversely, irregularly or spirally arranged, and they often
have anterior spines or projections or a posterior pedicel. The taxonomy of the loricate
choanoflagellates (Acanthoecidae) is based almost entirely on the structure of the lorica
determined by the arrangement of costae and the differentiation of costal strips. Within a
single species there can be considerable variation in cell dimensions (Thomsen et al., 1991a).

Loricate choanoflagelllates have been recorded frommany Antarctic localities (e.g. by Buck
& Garrison,1988; Marchant & Perrin, 1990). They are an abundant component of the
Antarctic marine plankton, comprising some 10-40% of the total number of living marine
protists (Marchant, 1985). They trap and ingest food, such as bacteria, cyanobacteria and
7
other particulate matter, and can occur in numbers in the order of 10 cells per litre, thus
suggesting they play an important role in Antarctic marine food webs.

Order Choanoflagellida

Family Codonosigidae

Monosiga ovata W.S.Kent (1880-82; pi. 2, figs 33-35)

Fig. 13.9a

ADDITIONAL REFERENCES: Leadbeater (1972d: 69, pi. 1), Hara et al. (1986: 72, figs 6, 7),
Marchant & Perrin (1986: fig. 3)

description: Cells solitary. Protoplast ovoid to ellipsoidal, 3.5-5 urn long, 2.8-3.2 urn
diameter. Collar of tentacles c. 6 urn long. Flagellum 8-1 urn long; short pedicel sometimes
1

present. Protoplast and base of tentacles surrounded by a delicate membrane sheath.

distribution: Indian sector of the Southern Ocean (Hara et al., 1986); Norway (Leadbeater,
1972d).

Family Acanthoecidae

Acanthocorbis tintinnabulum H.J. Marchant, in Marchant et al. (1987b: 18, figs 12-14)

Fig. 13.1a

description: Cells solitary. Lorica 11-12.5 urn long, 4.5 um diameter; single bell-shaped
lorica chamber 6.5 um long. Longitudinal costae 12-16, extending beyond chamber to form
spines 5-5.5 um long; anterior longitudinal costal strips (3.6-) 6-7 um long; costae taper
from c. 90-60 nm diameter towards anterior end on which is borne a claw; costal strips extend
c. 75% of their length beyond the lorica chamber; costal strips of lorica chamber c. 100 nm

*
Class Craspedophyceae or Class Choanoflagellida, or Superclass Choanoflagellata.

326
 CHOANOFLAGELLATES

diameter, c. 4 um long, with rounded ends. Transverse costae c. 40, spirally arranged;
distinctive anterior transverse costa present. Intact protoplasts have not been found.

distribution: Known only from Ellis Fjord, Vestfold Hills, East Antarctica (Marchant et ah,
1987b).

This species closely resembles A. unguiculata in having anterior spines with claw-bearing
tips and a similar arrangement of costae on the lorica chamber. However, A. tintinnabulum
differs from other Acanthocorbis species in the structure of the spines. In Acanthocorbis
apoda (B.Leadb.) S.Hara & E.Takah., A. unguiculata, and also Acanthoecopsis asymmetrica,
the spines are composed of at least 1.5 costal strips, whereas in A. tintinnabulum the spines
are less than a single costal strip in length.

Acanthocorbis unguiculata (H.A.Thomsen) S.Hara & E.Takah. (Hara & Takahashi,

1984: 827)

BASIONYM: Acanthoecopsis unguiculata H.A.Thomsen (1973: 22, figs 3-5; 1977: 93, figs 8-29)

SYNONYM: Pseudomicrosportella ornata Scagel & J. Stein (1961: 121 1, figs 9-1 1)

ADDITIONAL REFERENCES: van den Hoff & Franzmann (1986: 71), Marchant et al. (1987b: fig. 3)

Fig. 13.1b, c

description: Cells solitary. Lorica to 12 urn long, 11 um diameter; lorica chamber 7-8 urn
long, 9 um diameter. Longitudinal costae 13-15, extending as 5-6 um long spines beyond
the lorica chamber, in-curving, each bearing a small terminal claw. Transverse costae 6 or 7,
confined to posterior half of lorica. Protoplast spherical, 2.8-4.5 um diameter, located
posteriorly in lorica, enclosed by a sheath.

distribution: Inshore stations in Prydz Bay, East Antarctica (Marchant, 1985b); Greenland
(Thomsen, 1982).

Acanthocorbis unguiculata an important Antarctic heterotroph and has been observed to

is

ingest particles in um diameter and fluorescently labelled

the size range of 0.063-1.0
dextrans with molecular weight of 3,900-2,000,000 (Marchant & Scott, 1992).

Acanthoeca brevipoda W.N.Ellis (1929; 77, figs 6, 7)

ADDITIONAL REFERENCES: Leadbeater (1972e: 199, figs 7-10), Buck & Garrison (1983: 1264,
figs 18, 19)

Fig. 13. Id

description: Cells solitary. Lorica 9-10 um long, 3.8-4.6 um diameter. Longitudinal costae
10-12; anterior longitudinal costae protruding beyond lorica, forming spines 1.5-2 um long,
slightly incurved. Transverse costae c. 16-20, spirally arranged. Intact protoplasts not seen.

distribution: Ellis Fjord, Vestfold Hills, East Antarctica (Marchant et ah, 1987b); Weddell
Sea (Buck & Garrison, 1983); coastal Europe (Ellis, 1929; Leadbeater, 1972e; Thomsen,
1973; Leadbeater & Morton, 1974); North Pacific (Norris, 1965).

The Antarctic specimens have the same spiral arrangement of lorica costal strips as was
described in detail for coastal Denmark by Leadbeater (1972e). Smaller specimens, 6-7 um
long and with 8 longitudinal costae, were reported from the coastal North Pacific (Norris,
1965).

Acanthoeca spectabiiis W.N.Ellis (1929: 77, fig. 5)

ADDITIONAL REFERENCES: Hara & Takahashi (1984: 822, fig. 3A-I), Leadbeater (1979a: 62,
figs 13-36)

Fig. 13.2a

description: Cells solitary. Lorica (including the 12-17 anterior spines) to 18 um long, 4-5
um diameter; costal strips numerous, spirally arranged, converging at the posterior end of the

327
 CHOANOFLAGELLATES

lorica. Anterior spines sometimes curved inwards, comprising 3 costal strips, each 7-10 um
long. Protoplast obscured in anterior end of lorica chamber.

distribution: Coastal Antarctica, but not recorded from oceanic sampling stations;
Greenland (Leadbeater, 1979a); Japan (Hara & Takahashi, 1984).

In Antarctic specimens, the anterior spines sometimes curve inwards as in North American
material described by Norris (1965), but unlike the British specimens of Ellis (1929). Hara &
Takahashi (1984) described cells from Japan attached at their posterior end, or stalk, to
various substrata.

Acanthoecopsis asymmetrica H.A.Thomsen (1979: 21, figs 3, 22-25, 30, 31)

Fig. 13.2b

description: Cells solitary. Lorica c. 8 urn long, c. 7 um diameter at the level of the anterior
transverse costa; posterior lorica chamber 5.5 um long; longitudinal costae c. 15; transverse
costae 2. Arrangement of costal strips in posterior part of lorica complex and variable.
Longitudinal costae comprising 4 costal strips; anterior costal strips with rounded ends,
overlapping substantially with penultimate anterior costal strip as well as anterior transverse
costae; longitudinal costal strips somewhat thicker than transverse strips. Pedicel markedly
offset from long axis of lorica (hence the specific epithet). Protoplast ellipsoidal, to 4 um
long, 3 um diameter.

DISTRIBUTION: Occurs only rarely in the inshore waters of Prydz Bay, East Antarctica;
Finland (Thomsen, 1979).

The Antarctic organisms differ from the type description from Finland in that the cells are only
about half as long, and the anterior transverse costa is attached to the second longitudinal costal
strip (counted from the anterior end). Thomsen (1979) discussed the features distinguishing
Acanthoecopsis asymmetrica from Acanthoecopsis spiculifera R.E. Norris and Acanthocorbis
apoda.

Bicosta antennigera Moestrup (1979: 82, figs 41-44)

ADDITIONAL REFERENCES: Buck & Garrison (1983: 1264, fig. 26), Thomsen et al. (1990: 370, fig. 5)

Fig. 13.2c

description: Cells solitary. Lorica simple, Y-shaped, 25-60 um long, comprising 7 costal
strips, each to 8-10 um long. Longitudinal costae 2, with no obvious crossing-over, twisting
or spiralling, extending anteriorly to form 2 anterior spines to 37 um long, and posteriorly to
form a pedicel c. 25 um long, comprising a single costal strip. Transverse costae absent.
Protoplast ellipsoidal, 5-7 um long. Flagellum 16-18 um long.

distribution: Weddell Sea (Buck, 1981); New Zealand (Moestrup, 1979).

Bicosta antennigera shows no obvious twisting or spiralling of the longitudinal costae, a

feature that distinguishes it from B. spinifera. The pedicel, composed of a single costal strip,
also separates B. antennigera from B. spinifera and the closely allied B. minor (Reynolds)
B.Leadb. (Buck & Garrison, 1983, fig. 7), both of which have a pedicel comprising 4 costal
strips. Specimens examined by Manton et al. (1980) measured 40-80 um long. One small
specimen in the current study was 17 um long.

Bicosta spinifera (Throndsen) B.Leadb. (Leadbeater, 1978: 514)

BASIONYM: Salpingoeca spinifera Throndsen (1970b: 88)

ADDITIONAL REFERENCES: Manton et al. ( 980: 438, 1 pi. 3, 4), Thomsen et al. ( 1 990: 368, figs 2-4),
Thomsen & Larsen (1992: 55, figs 7-11)
Fig. 13. 2d

description: Cells solitary. Lorica slender, Y-shaped, (17-) 55 (-1 16) um long, with 7 costal
strips. Longitudinal costae 2, characteristically crossed-over in the anterior area of the

328
 CHOANOFLAGELLATES

Figure 13.1. Acanthocorbis and Acanthoeca species, a, Acanthocorbis tintinnabulum, SEM

(from Marchant et al, 1987). b, c, Acanthocorbis unguiculata, TEM (b) and SEM (c) (from
Marchant et al, 1987). d, Acanthoeca brevipoda, SEM Protoplast with single flagellum
surrounded by feeding tentacles. Scale bars: a-d = 2 urn.

329
 CHOANOFLAGELLATES

Figure 13.2. Acanthoeca, Acanthoecopsis and Bicosta species, a, Acanthoeca spectabilis.

TEM (from Marchant & Perrin, 1990). b, Acanthoecopsis asymmetrica, c. Bicosta antenn-
igera, SEM. d, Bicosta spinifera, TEM. Scale bars: a-d = 5 urn.

330
 CHOANOFLAGELLATES

Figure 13.3. Calliacantha, Cosmoeca and Crinolina species, a, Calliacantha natans, SEM.
b, Calliacantha simplex, SEM. c, Cosmoeca ventricosa, SEM. d, Crinolina aperta, TEM.
Scale bars: a-d = 5 urn.

33
 CHOANOFLAGELLATES

Figure 13.4. Diaphanoeca species, a. D. grandis. SEM (from Marchant & Perrin, 1986). b,
D. multiannulata, LM. c, D. pedicellata, TEM. d, D. sphaerica, TEM. Scale bars: a-d = 5 jam.

332
 CHOANOFLAGELLATES

protoplast, extending anteriorly to form 2 anterior spines and posteriorly to form a pedicel
comprising 4 costal strips. Transverse costae absent. Protoplast ellipsoidal, 5-9 pm long.

DISTRIBUTION: Abundant in Antarctic waters at temperatures below 10°C (Manton et al.,

1980; Buck, 1981; Thomsen & Larsen, 1990).

Some specimens have longitudinal costae that are obliquely arranged at their junction with
the posterior spine thus complicating the separation of B. spinifera, B. antennigera and
B. minor. See the discussion under Bicosta antennigera (above) as well as Thomsen et al.
(1990) and Hara & Takahashi (1987) for further discussion of B. minor.

Calliacantha natans (J.Grantved) B.Leadb. (Leadbeater, 1978: 514)

BASIONYM: Salpingoeca natans J.Grontved (1956: 9, fig. 6)

ADDITIONAL REFERENCES: Buck & Garrison (1983: 1264, fig. 29), Thomsen et al. (1990: 370,
figs 6-8, 17, 18)

Fig. 13.3a

description: Cells solitary. Lorica 32-35 pm long overall; lorica chamber 9-10 urn long.
Longitudinal costae 3; transverse costa 1. Anterior spines 3, 15 pm long, not continuous with
longitudinal costae; pedicel to 6 pm long. Protoplast ellipsoidal to obovoid, 5 pm long.

distribution: Common at inshore and oceanic stations of Prydz Bay, East Antarctica as well
as offshore in the vicinity of Casey Station; Weddell Sea (Buck & Garrison, 1983); larger
cells (to 60 pm long) reported from Lutzow-Holm Bay (Takahashi, 1981); Danish waters
(Grontved, 1956).

Calliacantha natans can be recognised by its 3 anterior spines which are not continuous with
the longitudinal costae (cf. C. simplex), and the relatively complex arrangement of costal
strips around the lorica chamber. Thomsen et al. (1990) reported "considerable variability in
lorica dimensions and proportions" for C. natans and C. simplex and presented data
illustrating the bimodal nature of size distributions of the two species. However, unlike
Arctic specimens, the size of Antarctic C. natans appears to be constant.

Calliacantha simplex Manton & Oates (1979a: 299, figs 1-30)

ADDITIONAL REFERENCES: Buck & Garrison (1983: 1264, fig. 28), Thomsen et al. (1990: 370,
figs 9, 10, 18, 19)

Fig. 13.3b

description: Cells solitary. Lorica c. 34 pm long overall; lorica chamber 1 pm diameter. 1

Longitudinal costae 4; transverse costae 2. Anterior spines 4, to 14 pm long, continuous with

longitudinal costae; pedicel sometimes present, to 8 pm long. Protoplast spherical, to 4.5 pm
diameter.

distribution: Common at inshore and oceanic stations of Prydz Bay, East Antarctica;
Weddell Sea (Buck, 1981); Alaska, Arctic Canada, Britain, the Galapagos Is. and South
Africa (see Manton & Oates, 1979b).

Cells were found in equal frequency with or without a pedicel, and the 2 transverse costal
stripswere obvious in both LM
and EM
preparations. Manton & Oates (1979a) reported
considerable variability in overall dimensions.

Cosmoeca norvegica H. A. Thomsen, in Thomsen & Boonruang (1984: 170, figs 11, 12)

ADDITIONAL REFERENCES: Moestrup (1979, as "Pleurasiga sp. N"), Thomsen et al. (1990: 371)

Fig. 13.9b

DESCRIPTION: [after Thomsen & Boonruang (1984)] Cells solitary. Lorica cup-shaped, 15-17 pm
long overall, 12.5-15 pm diameter at lorica end. Longitudinal costae 10; transverse costae 3;
the 2 anteriormost transverse costae each comprising 10 costal strips; the third less

333
 CHOANOFLAGELLATES

conspicuous and comprising 4-6 costal strips. Protoplast obovoid, 3-6 * 3-5 um (dried
specimen). Flagellum 15-25 um long.

distribution: Weddell Sea (Thomsen et al., 1991a).

Cosmoeca takahashii H. A. Thomsen, in Thomsen et al. (1990: 372, figs 22-26)

ADDITIONAL REFERENCE: Takahashi (1981: figs 17-19, as "Pleurasiga sp. 4")

Fig. 13.9d

DESCRIPTION: [after Thomsen et al. (1990)] Cells solitary. Lorica 16-22 um long overall, 15-19
um diameter at lorica end. Longitudinal costae 12; transverse costae 3. All costal strips are
thin rods; longitudinal costal strips project distally from the third anterior transverse costa,
forming a circle of spines. Protoplast obovoid, c. 4 x 7 um (dried specimen), in the centre of
the lorica. Flagellum to 25 um long.

distribution: Weddell Sea (Thomsen et al, 1991a).

Cosmoeca ventricosa H. A. Thomsen, in Thomsen & Boonruang (1984: 170, figs 14-17,
19-22)

ADDITIONAL REFERENCE: Thomsen et al. (1990: 371, figs 13-16, 20, 21)

Fig. 13.3c

DESCRIPTION: Cells solitary. Lorica barrel-shaped, to 31 um long and 22 um diameter.

Longitudinal costae 8; transverse costae 3. The 2 anteriormost transverse costae of equivalent
diameter, each comprising 8 costal strips; the third transverse costa much narrower,
comprising 3 or 4 costal strips. All costal strips are narrow rods. Protoplast obovoid, to 7.5
um long, situated in posterior end of the lorica.
distribution: Cells have been found only during the winter months in coastal Prydz Bay,
East Antarctica (Marchant & Perrin, 1990); Weddell Sea in autumn months (Buck &
Garrison, 1983).

This species is characterised by the absence of a pedicel and the occurrence of minute spines
projecting at the anterior end of the lorica.

Thomsen & Boonruang (1984) and Thomsen et al. (1990) documented a number of
morphological forms of this species. In Antarctic samples we have observed Thomsen and
Boonruang's "form A" (Fig. 13.3c) and "form B" (not illustrated).

Crinolina aperta (B.Leadb.) H. A. Thomsen (1976: 41)

BASIONYM: Diaphanoeca aperta B.Leadb., in Manton et al. (1975: 22, figs 25-32)

additional REFERENCES: Buck & Garrison (1983: 1264, fig. 32), Thomsen et al. (1990: 372, figs
27-32)

Fig. 13.3d

DESCRIPTION: Cells usually solitary, sometimes in clumps of 10-40 cells. Lorica barrel-shaped,
25^0 um long, 12-18 um diameter. Longitudinal costae 12-13, projecting beyond the
transverse costae at both ends, each comprising 8-9 costal strips; costal strips 4 um long, the
anteriormost strip thinner than the others. Costal strips of the 2 transverse costae L-shaped.
with the foot of the L pointing toward the anterior end of lorica, each 5 um long. Protoplast
ellipsoidal, 4 um diameter, situated centrally in the lorica.

distribution: Oceanic and inshore stations in Prydz Bay and Polar Aviation I., East
Antarctica; Weddell Sea (Buck, 1981; Thomsen et al., 1990).

There is considerable variation in the morphology of the costal strips. Details of lorica and
costal strip dimensions for material from the Weddell Sea were provided by Thomsen et al.
(1990).

334
 CHOANOFLAGELLATES

Figure 13.5. Kakoeca and Parvicorbicula species, a, K. antarctica, SEM. b, P. circularis,

SEM. c-e, P. socialis. c, LM Colony of cells, d, SEM Colony of cells, e, TEM Two
adjoining cells. Scale bars: a-e = 5 |um.

335
 CHOANOFLAGELLATES

Figure 13.6. Pleurasiga and Polyfibula species, a, Pleurasiga minima, TEM. b, Pleurasiga
orculaeformis,SEM. c, Pleurasiga reynoldsii. d, Polyfibula sphyrelata, TEM. Scale bars: a-d
= 5 (itn.

336
 CHOANOFLAGELLATES

Diaphanoeca grandis W.N.Ellis (1929: 74, fig. 10)

ADDITIONAL REFERENCES: Thomsen (1982: 6, figs 7, 8), Anderson (1988: 37)

Fig. 13.4a

DESCRIPTION: Cells solitary. Lorica 27-30 pm long, to 20 pm at posterior end. Longitudinal

costae 12, each comprising 7 costal strips, the 2 anteriormost strips extending 9-10 urn
beyond the anterior transverse costa. Transverse costae 4; anterior transverse costa composed
of a double row of costal strips. Protoplast ellipsoidal, c. 5 pm long, suspended centrally in
the lorica chamber, supporting a collar of c. 6 um long tentacles extending almost to the
level of the anterior transverse costa.

distribution: Inshore areas of Prydz Bay, East Antarctica (Marchant, 1985b); Greenland
(Thomsen, 1982); northern Pacific (Norris, 1965).

Some SEM, revealed a veil-like mesh on the inside of the

preparations, especially those for
lorica. This veil appears to shrink substantially during preparation for SEM leading to the
distortion of the lorica. As discussed by Marchant (1985b), the role of this structure has yet
to be determined. The chemical composition of the costal strips has been described in detail
by Thomsen (1982).

When present in high numbers, suspension feeders such as Diaphanoeca grandis and other
choanoflagellates may marine food webs. Ingestion experiments on
play a significant role in
D. grandis have shown of 0.063-1.0 um
that cells can ingest particles in the size range
diameter and fluorescently labelled dextrans with a molecular weight of 4,000-2,000,000
(Marchant & Scott, 1992), suggesting they are capable of exploiting a wide range of food
sources.

Diaphanoeca multiannulata K.R.Buck (1981: 47, figs 2 (holotype), 3-7)

ADDITIONAL REFERENCES: Buck (1980: 4, figs 3-12), Thomsen et al. (1990: 375, figs 37-48)

Fig. 13.4b

DESCRIPTION: Cells solitary. Lorica barrel-shaped, 50-75 um long (excluding pedicel), 25-35 um
diameter. Longitudinal costae 11-13, each comprising 8-9 costal strips, blunt at both ends,
extending c. 14 um beyond anterior transverse costa. Transverse costae 4 or 5, evenly spaced
throughout lorica length, each comprising 11-13 costal strips. Pedicel, when present, formed
by convergence of longitudinal costae, c. 15 um long, comprising 2 or 3 costal strips.
Protoplast spherical to obovoid, c. 6 um long, 3-4 um diameter; flagellum and collar of
tentacles arising from the anterior end; veil-like network of fine fibrillar material, suspended
from the anterior part of the lorica, often present.

distribution: Oceanic and inshore stations of Prydz Bay, East Antarctica; Antarctic waters
(Buck, 1980, 1981; Takahashi, 1981); Weddell Sea (Buck & Garrison, 1983).

The veil-like mesh on the inside of the lorica was discussed in Marchant (1985b). Prydz Bay
specimens of D. multiannulata may or may not have a pedicel. Loricae of Weddell Sea
material were reported by Buck (1981) to be 27-46 um long and 19-27 um diameter.

Diaphanoeca pedicellata B.Leadb. (Leadbeater, 1972e: 205, figs Id, 20, 21)

SYNONYM: Campanoeca pedicellata (B.Leadb.) Throndsen (1974: 1 10, fig. 30)

ADDITIONAL REFERENCES: Buck & Garrison (1983: 1264, fig. 30), Thomsen et al. (1990: 374,
figs 33-36)

Fig. 13.4c

description: Cells solitary or in colonies. Lorica to 45 um long (including pedicel); lorica

chamber 18-28 um long, 15 pm diameter; longitudinal costae 12-17; transverse costae 3.
Longitudinal costae each comprising 4-5 costal strips; anterior strip of each costa projects as
a spine beyond the anterior transverse costa. Costal strips of transverse costae thicker than

337
 CHOANOFLAGELLATES

those of longitudinal costae, or at least thickened in their middle. Pedicel, when present,
formed posteriorly from convergence of 7 longitudinal costae and c. 3 costal strips,
sometimes longer than lorica chamber. Protoplast obovoid, to 6 urn long, at posterior end of
lorica.

distribution: Oceanic and inshore sites of Prydz Bay, East Antarctica; deep (100 m) under-ice
stationsand surface open-water stations in the Weddell Sea (Buck & Garrison, 1983); North
Atlantic (Throndsen, 1974).

Diaphanoeca sphaerica H.A.Thomsen (1982: 20, figs 51, 52 (holotype), 53-58, 60, 63)

ADDITIONAL REFERENCE: Marchant & Perrin (1986: 209, fig. 5)

Fig. 13.4d

description: Cells solitary or Lorica bell-shaped, 22-30 urn long, to

in sphaerical colonies.
18 um diameter; longitudinal costae 13-15; costae 3. Longitudinal costae
transverse
comprising 9 or 10 costal strips; anterior costal strips thinner than those of the rest of the
lorica. Number of costal strips in each transverse costa equal to the number of longitudinal
costae. Arrangement of costal strips at the base of the lorica complex often appearing
stellate. Intact protoplasts not found in Prydz Bay material.

distribution: Burton Lake, a coastal lagoon in the Vestfold Hills, Prydz Bay, East
Antarctica (Marchant & Perrin, 1986); Disko Bay, West Greenland (Thomsen, 1982).
Although the Prydz Bay cells were solitary, Thomsen (1982) reported that cells in West
Greenland samples were "commonly united into sphaerical colonies (hence the specific
epithet) with the lorica apertures pointing towards a common cavity. Cells in the colonies are
arranged in clusters of six, and neighbouring cells are united along adjoining longitudinal
costae between the two anterior transverse costae". Thomsen also described protoplast
dimensions of c. 1 x 4 um.

Kakoeca antarctica K.R.Buck & H.J.Marchant, in Buck et al. (1990: 392, figs 1, 2, 3
(holotype), 4-10)

Fig. 13.5a

description: Cells solitary. Lorica barrel-shaped and tapering towards the posterior end, to
20-22 um long, 13-18 um diameter; oral diameter 4.8 um; longitudinal costae c. 20;
transverse costae 12; pedicel absent. Longitudinal and transverse costal strips c. 5 um long,
curved, all with simple ends except those at the distal ends of posterior longitudinal costal
strips which are dilated at their junctions. Longitudinal costae to the outside of transverse
costae. Anterior transverse costae spirally arranged; posterior transverse costal strips greater
in diameter than those at anterior end of lorica. Protoplast spherical, c. 6 um diameter,
supported by a solid suspensory membrane attached to posterior end of lorica; collar of
tentacles 4.5 um long. Flagellum 24 um long (including a 5 um long hairpoint), arising from
anterior end of protoplast.

distribution: Sea-ice near Davis Station, East Antarctica; Weddell Sea, in the sea-ice (Buck
etal, 1990).

The most conspicuous component of the lorica of this choanoflagellate. particularly if the
protoplast is absent, is the membranous sheath that supports the cell. This membrane has a
central hole and appears to be attached directly to the lorica at points along its anterior
margin and at its posterior apex.

Parvicorbicula circuiaris H.A.Thomsen (1976: 41, figs 24-28, 29 (holotype), 30-33)

ADDITIONAL REFERENCES: Buck & Garrison (1983: 1264, fig. 22), Thomsen et al. (1990: 378,
figs 49-57)

Fig. 13.5b

338
 CHOANOFLAGELLATES

description: Cells solitary. Lorica funnel-shaped, to 18-20 um long, 18-30 urn diameter;
longitudinal costae 4; transverse costae 2. Longitudinal costae each comprising 3 costal
strips; base of lorica chamber constructed of 4-6 costal strips, of which 4 are an extension of
the longitudinal costae; posterior costal strip abuts the second most posterior strip for c. 25-
33% of their length. Anteriormost transverse costa comprising 8 costal strips; anterior tips of
the anterior costal strips swollen, bifurcated and joined at the centre of every other strip in
the anterior transverse costa, these strips abut every other intersection between strips in the
posterior transverse costa. Some specimens with other transverse costae comprised of 6 or 7
costal strips. Protoplast obovoid, 10 urn long, 5 um diameter, located in base of lorica
chamber, suspended by a sheath from the second most posterior costal strip.

distribution: Inshore and oceanic stations of Prydz Bay, East Antarctica; Liitzow-Holm Bay
(Takahashi, 1981, as "species 9"), Weddell Sea (Buck & Garrison, 1983); Denmark
(Thomsen, 1976).

While there are differences between the Danish type material of P. circularis and material
collected in Antarctica, we do not consider the differences significant enough to warrant the
recognition of a new species at this time. The type specimen of P. circularis has 8 costal
strips in the anterior costae and only 6 in the posterior costae. Prydz Bay specimens are
similar in costal strip morphology to those from Liitzow-Holm Bay and the Weddell Sea.
Thomsen (1976) discussed the differences between P. circularis and the closely allied
P. quadricostata Throndsen from Denmark.

Parvicorbicula ongulensis E.Takah. (Takahashi, 1981: 7, figs 13, 14)

Fig. 13.9c

description: Cells solitary. Lorica funnel-shaped, 34-35 um long (including pedicel), c. 15

urn diameter; longitudinal costae 14; transverse costae 3. Longitudinal costae each
comprising 5 costal strips 5-5.5 um long, converging at the posterior end to form the
pedicel; anteriorly, 1 costal strip of each costa projects beyond the anterior transverse costa;
pedicel comprising 2 or 3 costal strips 10-10.5 um long. Anterior and second transverse
costae comprising 14 and 15 costal strips, respectively; costal strips 4-4.2 um long, slightly
curved, slightly thicker than the longitudinal costal strips but taperingboth ends.
at
Protoplast ovoid, 3.5 um long, 3 um diameter, located posteriorly in lorica. Flagellum 8 um
long.

distribution: Only known from under sea-ice in Liitzow-Holm Bay, East Antarctica
(Takahashi, 1981).

Parvicorbicula socialis (Meunier) Deflandre (1960: 183)

basionym: Corbicula socialis Meunier (1910: 79)

ADDITIONAL REFERENCES: Manton et al. (1975: 1935, figs 1-4, 11-38), Buck & Garrison (1983:
1264, fig. 21), Hara & Tanoue (1984b: 6), Thomsen et al. (1990: 380, figs 59-65)

Fig. 13.5c-e

description: Cells solitary or forming extensive colonies of cells adhering together at their
anterior costae. Lorica funnel-shaped, 11-13 urn long, 16-19 um diameter at anterior edge;
longitudinal costae 10; transverse costae 2; with or without a pedicel. Longitudinal costae
comprising 10 costal strips, 6 um long, rounded and flattened, forming characteristic T-
junctions where they meet the anterior costa. Transverse costae comprising 10 costal strips,
c. 5 jam long. Protoplast ellipsoidal, 3.5-4 um diameter.

distribution: Widely distributed in inshore and oceanic Antarctic waters and in many other
parts of theworld (Hara & Tanoue, 1984), in water temperatures of-2.4°C (Buck, 1981) to
25°C (Thomsen, 1978a).
Cell size ranges have been documented for Antarctic specimens (Buck, 1981; Takahashi,
1981; Thomsen et al., 1990) as well as for Arctic 'summer' specimens (Manton et al.,

339
 CHOANOFLAGELLATES

1976a). Additional illustrations and descriptive details of Antarctic specimens were provided
by Hara & Tanoue (1984) and Thomsen et al. (1990).

Pleurasiga minima Throndsen (1970a: 109, figs 21-25)

additional references: Buck & Garrison (1983: 1264, fig. 23), Thomsen et al. (1990: 380,
figs 66, 67)

Fig. 13.6a

description: Cells solitary. Lorica barrel-shaped, 10-14 urn long, 10-12 um diameter;
longitudinal costae 7; transverse costae 2. Longitudinal costae comprising 3 costal strips,
with characteristic T-junctions where they meet the anterior transverse costa. Transverse
costae comprising 7 costal strips. Protoplast obovoid, 4-6 um long, lying in close connection
with the basal part of the lorica; collar of tentacles extending to the anterior costa. Flagellum
c. 20 um long.

distribution: Sparse at oceanic and inshore stations of Prydz Bay, East Antarctica; Weddell
Sea (Buck & Garrison, 1983); North Atlantic (Throndsen, 1974); Californian coast (Thomsen
et al, 1991a).

Pleurasiga minima is essentially distinguished from its close ally P. reynoldsii by the latter
being conspicuously larger in size and having longitudinal costae that merge into pairs where
the penultimate and ultimate longitudinal costal strips meet, resulting in only 4 longitudinal
costae at the posterior end of the cell (Thomsen et al., 1990). Cells intermediate in size
between the two species have also been found at oceanic sites in the vicinity of Casey
Station, East Antarctica.

Pleurasiga orculaeformis J.Schiller (1925b: pi. 4, fig. 24)

ADDITIONAL REFERENCES: Leadbeater (1973: 237, figs la, b, 2b, pi. 14a-d, pi. 15a, b),
Hallegraeff (1983: 501, fig. 37); Buck & Garrison (1983: 1273, fig. 24)

Fig. 13.6b

description: Cells solitary. Lorica barrel-shaped, 25-28 um long, 16-21 um diameter;

longitudinal costae 9-13; transverse costae 3. Longitudinal costae comprising 4 costal strips;
in some specimens the anterior costal strips extend c. 25% of their length beyond the anterior
transverse costa. Protoplast subspherical, 3 um diameter, lying in close connection with the
base of the lorica.

distribution: Inshore and oceanic sites in Prydz Bay, East Antarctica; coast near Sydney,
Australia, coastal waters, New Zealand and Thailand (Hallaegraaf, 1983); Adriatic and Bay
of Algiers (Leadbeater, 1974); Disko Bay, West Greenland (Thomsen, 1982).

Details of the lorica and inner loricate collar were discussed by Leadbeater (1973).
Takahashi (1981) illustrated this organism (as "species 4") with longitudinal costal strips
extending beyond the anterior transverse costa, as did Hallegraeff (1983) in material from
the East Australian Current. Specimens with the longitudinal costae terminating at the
anterior transverse costa were reported by Buck & Garrison (1983).

Pleurasiga reynoldsii Throndsen (1970a: 106, figs 11 (holotype), 12-20)

ADDITIONAL REFERENCE: Manton et al. (1976a: 1940, figs 9, 10, 61-71)

Fig. 13.6c

description: Cells solitary. Lorica funnel-shaped, to 23 um long, 16-22 um diameter;

longitudinal costae 7; transverse costae 2. Longitudinal costae comprising costal strips
sometimes with slightly flattened bifurcate ends; anterior ends of longitudinal costae forming
T-junctions with the anterior transverse costa; 6 of the 7 longitudinal costae merge in pairs at
the level of the protoplast. Protoplast slightly obovoid, 7-8 um long, lying close to the base
of the lorica chamber, flanked by 4 costal strips. Flagellum extending beyond the anterior
costa.

340
 CHOANOFLAGELLATES

Figure 13.7. Saepicula and Spiraloecion species, a, b, Saepicula leadbeateri, TEM. c, d,

Spiraloecion didymocostatum, TEM (from Marchant & Perrin, 1986). Scale bars: a-d = 5 um

341
 CHOANOFLAGELLATES

Figure 13.8. Stephanoeca species, a, S.complexa, SEM (from Marchant et a/., 1987). b. c,
S. diplocostata var. paucicostata, TEM (b) and SEM (c) (from Marchant et ai. 1987). d,
S. norrisii, SEM (from Marchant et ai, 1987). Scale bars: a-d = 5 ^m.

342
 CHOANOFLAGELLATES

distribution: Inshore and oceanic areas off the Vestfold Hills, Prydz Bay, East Antarctica;
Arctic waters (Throndsen, 1970); widespread throughout the Northern Hemisphere (see
Leadbeater, 1973).

Cells intermediate in size between Pleurasiga reynoldsii and P. minima have also been found
in Prydz Bay (see discussion under P. minima).

Polyfibula sphyrelata (H.A.Thomsen) Manton, in Manton & Bremer (1981: 290,

figs 1-5)

BASIONYM: Pleurasiga sphyrelata H.A.Thomsen (1973: 24, figs 22-26)

SYNONYM: Campanoeca dilatata Throndsen (1974: 109, figs 2, 8, 29)

ADDITIONAL REFERENCE: Thomsen el al. (1990: 382, figs 68, 69)

Fig. 13. 6d

DESCRIPTION: Cells solitary. Lorica barrel-shaped, 15-20 jam long, c. 10 jam diameter; longit-
udinal costae 7 or 8; transverse costae 3. Longitudinal costae extend for about 10% of their
length beyond the anterior transverse costa, each comprising 4 costal strips. Transverse costae
comprising 7 or 8 costal strips; costal strips of anterior and middle costae, but only one of the
posterior costa, bearing a small square facet in the middle of the costal strip. Protoplast
ellipsoidal, 4 um long, located posteriorly in the lorica; collar of tentacles c. 3 urn long.

DISTRIBUTION: Inshore areas of Prydz Bay, East Antarctica; New Zaland (Moestrup, 1979);
South Africa (Manton et al., 1975); type locality, Denmark (Thomsen, 1973).

Posteriorly, this organism resembles Polyfibula elatensis (H.A.Thomsen) Manton and

P. caudata (B.Leadb.) Manton. However, the arrangement of the remainder of the lorica,
especially insofar as the anterior ends of the longitudinal costae neither make T-junctions (as
in P. caudata), nor that these costal strips extend well beyond the anterior transverse costal
strips as spines (as in P. elatensis), indicates that this organism should be regarded as
P. sphyrelata. In the Prydz Bay material, the square costal strip facets were not well
developed on the middle and posterior transverse costae.

Saepicula leadbeateri E.Takah. (Takahashi, 1981a: 9, fig. 20)

ADDITIONAL REFERENCES: Marchant & Perrin (1986: 209, fig. 4), Thomsen et al. (1990: 382, fig. 70)

Fig. 13.7a, b

DESCRIPTION: Cells solitary. Lorica bell-shaped, to 16.5 um long, 8 um diameter at the level
of the anterior transverse costa. Longitudinal costae 9 or 10, longitudinally or obliquely
arranged, each comprising 3 thick costal strips; anterior costal strips 7.2 jam long, 0.1 um
thick, sharply pointed, extending beyond the transverse costa; centrally located costal strips
blunt-ended; posterior costal strips variable in thickness, but thicker than the others.
Transverse costa solitary, comprising c. 20 thick round-ended costal strips, located
c. halfway along the length of the anterior longitudinal costal strip; costal strips 3 |am long,

0.3 um thick. Pedicel, if present, variable in length but shorter than the lorica. Intact
protoplasts not seen in the Prydz Bay material.

DISTRIBUTION: Sea-ice, Burton Lake (tidal lagoon in the Vestfold Hills), Crooked Fjord and
other coastal waters off Davis Station, East Antarctica (Marchant & Perrin, 1986); Liitzow-
Holm Bay (Takahashi, 1981a).
There appears to be some variation in the thickness of the costae in the posterior of the
lorica. In organisms with a long pedicel the costae are thin, some with pointed ends similar
to the costal strips of the anterior longitudinal costae. Organisms with a short pedicel
sometimes have markedly thickened, posterior costal strips. Specimens with either long or
short pedicels were observed in Prydz Bay (Marchant & Perrin, 1986).

343
 CHOANOFLAGELLATES

Spiraloecion didymocostatum H.J.Marchant & R.Perrin (1986: 210, figs 6, 7)

ADDITIONAL REFERENCE: Takahashi (1981a: 9, pi. 6, fig. 21, as Sp. no. 15)'

Fig. 13.7c, d

description: Cells solitary. Lorica conical, c. 17 (jm long including the pedicel, 8 um
diameter. Lorica chamber c. 10 um long. Longitudinal costae 7-11, spirally arranged at the
posterior end of lorica, interwoven with a variable in number of transverse costae; anterior
transverse costa joins the longitudinal costae at the junction of the first and second costal
strips of the longitudinal costae; all costal strips of the lorica chamber 3.2 um long. Pedicel
to 15 um long, comprising thickened apparently flattened costal strips 1.5-3.6 um long and
c. 0.13 um thick that also form part of the posterior end of the lorica chamber. Intact

protoplasts not seen.

Figure 13.9. Monosiga, Cosmoeca, Parvicorbicula and Stephanoeca species, a, M. ovata,

SEM Cell with tentacles (arrow) (from Marchant & Perrin, 1986). b, C. norvegica, TEM
(from Thomsen & Boonruang, 1984). c, P. ongulensis, TEM (from Takahashi, 1981). d,
C. takahashi /. TEM (from Thomsen et al, 1990). e, f, 5. cupula, TEM (from Leadbeater,
1972). Scale bars: a. e, f, =
1 urn; b-d = 5 um.

344
 CHOANOFLAGELLATES

DISTRIBUTION: Burton Lake (tidal lagoon in the Vestfold Hills), East Antarctica (Marchant &
Perrin, 1986); at the bottom of the sea-ice in Liitzow-Holm Bay (Takahashi, 1981a, as Sp.
no. 15?.

Marchant & Perrin (1986) noted that Spiraloecion superficially resembles some species of
Acanthoecopsis. However, the number of costae and details of costal strip arrangement and
morphology separate the two genera.

Stephanoeca complexa (R.E.Norris) Throndsen (1974: 1 18, fig. 45)

BASIONYM: Pleurasiga complexa R.E.Norris (1965: 599, fig. 21)

ADDITIONAL REFERENCES: Marchant & Perrin (1986: 208, fig. 2), Marchant et al. (1987b: 15, figs
7,8)

Fig. 13.8a

description: Lorica somewhat waisted to form anterior and posterior chambers, 13.5-15 um
long excluding pedicel (when present); anterior chamber 8-11 p.m diameter; posterior
chamber 4-6.5 um diameter; longitudinal costae 10-12; transverse costae variable, 4 or 5 in
basal part of lorica. Arrangement of costal strips irregular and rather complex; longitudinal
costae obliquely orientated, some of the costal strips of the posterior chamber thickened.
Protoplast ovoid, to 6.5 um long, 2.2-5 um diameter

distribution: Ellis Fjord, Vestfold Hills, East Antarctica (Marchant & Perrin, 1986); North
Atlantic (Throndsen, 1974); north-east Pacific (Norris, 1965).

This species is readily recognised by its waisted outline and the obliquely orientated costae.
Clusters of Stephanoeca complexa cells are sometimes found associated with other
organisms, especially flagellates.

Stephanoeca cupula (B.Leadb.) H.A.Thomsen (1988: 320, figs 16-22)

BASIONYM: Pleurasiga cupula B.Leadb. (Leadbeater, 1972b: 201, figs le (holotype), 16, 17)

ADDITIONAL REFERENCE: Hara et al. (1986: 75, fig. 12)

Fig. 13.9e, f

DESCRIPTION: Cells solitary. Lorica cup-shaped, somewhat waisted to form two chambers,
6-10 um long, 6-9 p.m diameter; longitudinal costae 8-11; transverse costae 3 in anterior
chamber, 2 in posterior chamber. Longitudinal costae each comprising 3 costal strips, the
posteriormost converging spirally with several transverse costal strips towards the posterior
end of the lorica. Protoplast ovoid, 2-3 um long, 1-2 um diameter, located in posterior
chamber; flagellum 4-5 um long; collar of 10-15 tentacles, each c. 2 u.m long.

DISTRIBUTION: Indian sector of the Southern Ocean (Hara et al., 1986); coastal Denmark
(Leadbeater, 1972b; Thomsen, 1973).

Further details of lorica strucure were described and illustrated by Thomsen (1988).

Stephanoeca diplocostata W.N.Ellis var. paucicostata Throndsen (1969: 178, fig. 14)

ADDITIONAL REFERENCES: Marchant et al. (1987b: 17, figs 9, 10), Marchant & Perrin (1990: 504)

Fig. 13.8b, c

description: Cells solitary. Lorica somewhat waisted to form anterior and posterior
chambers, 16-17 (im long overall; anterior chamber 10 um long, 9 um diameter; posterior
chamber 6.5 um long, 6.4 um diameter; longitudinal costae c. 13; transverse costae 3 in
anterior chamber, 4 in posterior chamber; pedicel sometimes present. Longitudinal costae each
comprising 5 costal strips; anteriormost costal strips forming T-junctions or short stalks where
they meet the anterior transverse costa. Transverse costal arrangement difficult to discern; a
double row of transverse costae surrounds the middle of the anterior lorica chamber [as
described by Throndsen (1969)]; another costa at the waist of the lorica; 4 costae in the

345
 CHOANOFLAGELLATES

posterior chamber. Protoplast ellipsoidal to obovoid, 4.2 um long, situated in the posterior
lorica chamber, with a collar of tentacles and a flagellum in the anterior chamber.

distribution: Inshore Prydz Bay, East Antarctica; North Atlantic waters (Throndsen, 1969,
1974).

Stephanoeca norrisii H.A.Thomsen (1973: 10, figs 15, 16)

SYNONYM: Sportelloeca kentii (W.N.Ellis) R.E.Norris (1965: 602, figs 20, 24)

ADDITIONAL REFERENCE: Marchant et al. (1987b: 18, fig. 11)

Fig. 13. 8d

description: Cells solitary. Lorica to 23 urn long (including pedicel, if present), 8 \im
diameter; lorica chamber 15-18 um long; costal strip arrangement distinctive. Costae at
anterior and posterior ends of lorica transversely arranged and obliquely orientated at their
extremities; median costal strips longitudinal, c. 80, closely packed; costal strips of posterior
part of lorica thicker than others. Protoplast spherical, c. 2.6 um diameter, located just below
centre of lorica. Flagellum extending beyond the lorica, terminating in a hair-point c. 25% of
total flagellar length.

distribution: Ellis Fjord, Vestfold Hills, East Antarctica (Marchant et al., 1987b); Danish
waters (Thomsen, 1973); North Pacific (Norris, 1965, as Sportelloeca kentii).

Norris's (1965) Sportelloeca kentii was based on Stephanoeca kenti W.N.Ellis (1929: 73),
but these descriptions have since been recognised as representing two distinct species.
The name Stephanoeca kenti is retained for a choanoflagellate without longitudinal costae
(Ellis, 1929, Sportelloeca kentii (with a distinct median band of longitudinal costae)
fig. 1 1);
was returned to Stephanoeca by Thomsen (1973) who honoured Norris with the epithet.

346
 14. CILIATES
Wolfgang Petz

Ciliates are a well-defined, monophyletic group of unicellular protists (protozoa). With the
exception of the 'primitive' karyorelictids, they are characterised by having two types of
nuclei, i.e. at least one macronucleus and one micronucleus. Another distinctive feature is
conjugation, a sexual phenomenon during which gamonts partially and temporarily fuse
(gamontogamy). On the outside, the ciliate cell is covered by few to many cilia or compound
ciliary organelles (cirri) anchored in the cortex (outer cell layer) by a complex infraciliature
(kinetosomes and closely associated microtubules and microfibrils). Somatic and oral cilia
and cirri, which are mainly used in locomotion and food gathering, are often arranged in
rows (kineties), but these can be considerably reduced or even lacking in some groups or in
some stages of the life cycle. Many ciliates are mobile throughout their entire life cycle, but
some (e.g. folliculinids, suctorians and most peritrichs) are sessile in their adult stage.

Ciliates consume a wide range of food, e.g. bacteria, cyanobacteria, microfungi, microalgae
(often diatoms) and other protists (including ciliates), small metazoans, organic particles and
dissolved organic matter. Several taxa (usually oligotrichs) are mixotrophic, and at least one
species (Myrionecta rubra) is apparently exclusively autotrophic due to a cryptophycean
endosymbiont (Lindholm, 1985).

Most freshwater and marine habitats under both

ciliate species are free-living in terrestrial,
oxic and anoxic conditions, some endocommensals, a few are true parasites that
are ecto- and
can cause considerable economic damage in fish farms, and very few are pathogens. Some
species can occur in enormous abundance in the plankton and form blooms ('red tides'). The
ciliates comprise several size classes, i.e. the smallest forms are less than 20 urn long (e.g.
mesodiniids) whereas the largest representatives hitherto found in the Southern Ocean
(Condylostoma sp.) measured almost 2000 [im in length (Petz et ai, 1995). However, most
species belong to the microplankton size fraction (20-200 urn).

Loricate (mainly tintinnids) and aloricate ciliates are an important component of marine
planktonic and benthic food webs. Within the heterotrophs, they are among the numerically
dominant groups (Sorokin, 1999). In the plankton of the Southern Ocean, ciliates can reach
abundances of up to 29,000 individuals per litre (Sorokin, 1999) and account for 54-64% of
the total zooplankton production (Sushin et al., 1986). Associated with a Phaeocystis bloom,
densities may be as high as 150,000 individuals per litre (Davidson & Marchant, 1992b). As
in other oceans, aloricate species are usually more abundant than tintinnids (Garrison &
Gowing, 1993). High numbers of aloricate ciliates occur also in the endopagial (brine-
channels) of Antarctic sea-ice (Garrison, 1991b; Petz et al., 1995) where they, along with
aplastidic flagellates, form the dominant portion of the heterotrophic biomass (Garrison &
Buck, 1991). A recent investigation in the Ross Sea showed, for instance, that the ciliate
population can exceed 400,000 individuals per litre of melted ice (biomass 2.8 mg carbon per
litre) in older brown ice (Petz, unpublished data).

Ciliates are identified by body shape, size and colour, the shape and arrangement of the
somatic and oral ciliature, the shape and number of macro- and micronuclei, the position and
number of contractile vacuoles, and other characters in certain groups (e.g. extrusomes,
nematodesmata, loricae, body appendages, dorsal silverline system, behaviour). Not all of
these features are recognisable in fixed specimens, so that in vivo observations are frequently
required for an accurate identification. Tintinnid loricae exhibit considerable infraspecific
variability between generations in many (all?) species and are consequently a rather
unreliable taxonomic character (e.g. Laval-Peuto, 1981; Van der Spoel, 1986; Petz et ai,
1995; Wasik, 1998).

Phylum Ciliophora

347
 CILIATES

Besides marine biotopes of the Southern Ocean, ciliates are also widespread in soil, mosses,
freshwater lakes and meltpools in Antarctica, although they often occur only in low
concentrations (e.g. Thomas, 1965; James et al, 1995, 1998; Laybourn-Parry et al, 1997;
Petz, 1997, 2003; Petz & Foissner, 1997; Valbonesi & Petz, 1998).

This compilation illustrates the considerable diversity of ciliate species in the Southern
Ocean. At least 161 species, 47 tintinnid and 114 aloricate taxa from 85 genera, have been
recorded and are described here; several others were only incompletely identified. In
addition to the genera treated below, one or more unidentified species of Acineta,
Chilodonella, Coleps, Dileptus, Ephelota (epizoic on krill), Epiclintes, Metacystis,
Paraurostyla, Parundella, Peritromus, Rhabdoaskenasia, Scyphidia, Trochilia and
undetermined, peritrich ciliates (Fig. 14.193) were also found in the Southern Ocean
(Thompson, 1972; Thompson & Croom, 1978; Corliss & Snyder, 1986; Heinbokel & Coats,
1986; Rakusa-Suszczewski & Nemoto, 1989; Valbonesi & Luporini, 1990c; Petz et al, 1995;
Alder & Thompson, 2000; Song & Wilbert, 2002a; Stankovic et al, 2002; Petz, unpublished
data; Petz & Valbonesi, unpublished data; Scott, unpublished data). This suggests that ciliate
species diversity is even higher than is currently known.

Eufolliculina ampulacea Hadzi (1951: 16, fig. 1) was established for a population of
Folliculina ampulla (Muller) Lam. found in the Mediterranean Sea and described by
Laackmann (1910b: fig. 10) in a report on Antarctic ciliates. This species was included in error
in the list of Antarctic folliculinids by Ringuelet (1957) and, therefore, is not documented here.

CLASS HETEROTRICHEA
Order Heterotrichida

Family Folliculinidae

Ascobius simplex (Dons) Hadzi (1951: 119)

BASIONYM: Semifolliculina simplex Dons (1918: 78, fig. 2)

SYNONYMS: Lagotia simplex (Dons) Dons (1928: 7, fig. 7)

Folliculina simplex (Dons) Kahl (1932: 473, fig. 77.23, 26)

ADDITIONAL REFERENCES: Faure-Fremiet (1936: 1153), Das (1948: 452), Ringuelet (1953a: 196;
1953b: 231)

Fig. 14.1a, b

DESCRIPTION: Lorica 135-200 * 65-120 urn, ovoid, dark green, flattened; neck very short;
opening narrow; ampulla lying (attached to substratum by its broad lateral side). Trophont
190-220 urn long, coloured blue due to pigment granules; short peristomial wings rounded,
unequal. Macronucleus 1. Epibiotic on algae, molluscs and isopods.

distribution: Bellingshausen Sea (Ringuelet, 1953a); Atlantic (Dons, 1918; Faure-Fremiet,

1936; Das, 1948).

A new variety (now subspecies) Ascobius simplex Irregularis was described by Ringuelet
(1953a: 198, figs 4-7) for Antarctic specimens with an irregular lorica outline.

Folliculina ampulla (Muller) Lamarck (1816: 30)

BASIONYM: Vorticella ampulla Muller (1786: 283, pi. 40, figs 4-7)

See Hadzi (1951) for synonymy.

ADDITIONAL REFERENCES: Kahl (1932: 470), Hadzi (1951: 111)

Fig. 14.2

description: Lorica 135-200 * 90-100 um, ovoid to broadly bottle-shaped, colourless;

348
 CILIATES

Figures 14.1-6. Ascobius, Folliculina and Lagotia species. 14.1, A. simplex, broad side from
life (a, from Faure-Fremiet, 1936), and A. simplex irregularis, after fixation (b, broad side,
from Ringuelet, 1953a). 14.2, F. ampulla, broad side from life (from Kahl, 1932). 14.3, L.
expansa, lateral side from life (from Mulisch et al., 1986). 14.4, L. faurefremieti, lateral side
from life (from Faure-Fremiet, 1936). 14.5, L. gigantea, lateral side after fixation (from
Kahl, 1932). 14.6, L. similis, lateral side after fixation (from Kahl, 1932). Scale bars: 1, 2 =
100 urn; 3, 5 = 150 u.m; 4, 6 = 200 um. Abbreviations: A, ampulla; C, collar; L, lorica; Mn,
macronucleus; N, neck; P, peristomial wing.

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 CILIATES

Figures 14.7-11. Parafolliculina, Pseudofolliculina and Heterostentor species. 14.7,

Parafollieulina amphora, broad (a) and narrow (c) sides from life (trophont contracted; from
Hadzi, 1951) and broad side after fixation (b, from Dons, 1914). 14.8, Parafolliculina
patagonica, broad side after fixation (trophont contracted; from Ringuelet, 1957). 14.9,
Parafolliculina violacea, broad (a) and narrow (b, trophont contracted) sides from life (from
Faure-Fremiet, 1936). 14.10, Pseudofolliculina melitta, lateral side after Fixation (trophont
contracted; from Laackmann, 1910b). 14.11, Heterostentor coeruleus, lateral side from life
(a) and infraciliature of anterior ventrolateral side (b) after protargol impregnation (from
Song & Wilbert, 2002). Scale bars: 7 = 50 urn; 8 = 80 urn; 9, 10 = 100 |im; 11a = 60 um.
Abbreviations: AZM, adoral zone of membranelles; B. base; M, closing membrane; S, stalk;
Sk, somatic kinety; Su, suture.

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 CILIATES

ampulla lying; neck rather short, rising at an angle of c. 45-60° from substratum; opening
slightly everted. Trophont dark green, contractile; short peristomial wings roundish to
elongate, dorsally scarcely separate. Macronucleus 1. Epizoic on bryozoans.

distribution: Indian sector of the Southern Ocean (Schroder, 1906a); North Atlantic
(Claparde & Lachmann, 1858); Greenland (V anhoffen, 1897); North Sea (Kahl, 1932);
White Sea (Mereschkowsky, 1879); widespread.

According to Hadzi (1951), several species are aggregated under the name F. ampulla. Thus,
for example F. ampulla sensu Laackmann (1910b) in the Southern Ocean is Lagotia gigantea
(see below).

Lagotia expansa (Levinsen) Hadzi (1951: 48, figs 6, 7)

BASIONYM: Filellum(?) expansion Levinsen (1893: 172, pi. 7)

SYNONYM: Folliculina expansa (Levinsen) Kramp (1911: 374)

ADDITIONAL REFERENCES: Dons (1911: 81), Hadzi (1951: 48), Ringuelet (1953a: 201; 1953b:
235), Mulisch e? a/. (1986:4)

Fig. 14.3

description: Lorica 270-730 x 120-220 pm, light blue-green, variable in shape; ampulla
lying, broad (120-380 x 120-220 pm); neck annulated (100-400 urn long), longitudinally
striated; opening everted. Trophont blue-green, contractile; peristomial wings equal, to 400
long and 55 urn wide. Macronucleus ellipsoidal. Epizoic on bryozoans, molluscs and
isopods; often in dense colonies.

distribution: Bellingshausen Sea, Atlantic, Pacific (Ringuelet, 1953a); Mediterranean Sea

(Hadzi, 1951).

Lagotia faurefremieti Hadzi (1951: 47)

SYNONYM: Folliculina aculeata Claparde & Lachmann sensu Faure-Fremiet (1936: 1 148, fig. 8)

ADDITIONAL REFERENCES: Ringuelet (1953a: 202), Mulisch et al. (1986: 4)

Fig. 14.4

DESCRIPTION: Lorica to more than 1000 um long, dark blue-green; ampulla lying,
comparatively slim (450-650 * 200-250 um); neck annulated (175-400 x 100 um); opening
everted. Trophont dark blue-green, but often appearing red under low magnification;
peristomial wings equal, c. 200 x 40 pm. Somatic kineties 70-80 longitudinal. Macronucleus
ellipsoidal. Epibiotic on bryozoans and red algae.

distribution: Bellingshausen Sea (Ringuelet, 1953a); Atlantic (Mulisch et al, 1986).

Lagotia gigantea (Dons) Dons (1934: 32)

BASIONYM: Semifolliculina gigantea Dons (1914: 67, figs A, B, pi. 1, figs 1-9)

SYNONYMS: Folliculina ampulla (Miiller) sensu Laackmann (1910b: 84, figs 16-25)

Folliculina gigantea (Dons) Kahl (1932: 472, fig. 77.9)

ADDITIONAL REFERENCES: Hadzi (1951: 81; L. gigantea considered to be possibly synonymous

with L. expansa)

Fig. 14.5

DESCRIPTION: Lorica 760-1870 x 100-250 pm, rather variable in size, brown to blue-green;
ampulla lying; neck sometimes rising at 70-90° from substratum, spirally annulated, often
longitudinally striated; opening usually distinctly everted (160-340 pm across), sometimes
with a second collar. Macronucleus globular. Epizoic on bryozoans.

distribution: Indian sector of the Southern Ocean (Laackmann, 1910b); North Atlantic
(Kahl, 1932).

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 CILIATES

Lagotia similis (Dons) Dons (1934: 32)

BASIONYM: Semifolliculina similis Dons (1915: 149, fig. 5)

SYNONYM: Folliculina similis (Dons) Kahl (1932: 471, fig. 77.3)

ADDITIONAL REFERENCES: Dons (1918: 77), Hadzi (1951: 85)

Fig. 14.6

description: Lorica 700 um long, flask-shaped, brown; ampulla lying; neck slightly
c.

annulated, rising at 90° from substratum; opening with a small collar. Trophont to 1000
c.

um long, green to brown; peristomial wings equal, elongate, each with a distal style-like
projection. Macronucleus 1, ellipsoidal to spherical. Epizoic on bryozoans.

distribution: Indian sector of the Southern Ocean (Dons, 1915).

Parafolliculina amphora Dons (1914: 83, figs C, D; pi. 2, figs 1-6)

ADDITIONAL REFERENCES: Faure-Fremiet (1936: 1161), Dons (1948: 7), Hadzi (1951: 172),
Ringuelet (1953b: 242; 1955: 180; 1957: 214)

Fig. 14.7a-c

description: Lorica 120-200 * 65-120 um, sac-like to flask-shaped, with a flattened

swollen ring and a distinct constriction in the anterior part (neck), blue, green or brownish
yellow to colourless; ampulla and part of neck lying; opening bent upwards, only indistinctly
widened, with a bipartite closing membrane (lid or flap) between ampulla and neck.
Trophont concolorous with the lorica; peristomial wings equal, distally pointed.
Macronucleus 1. Epibiotic, e.g. on algae, bryozoans and molluscs.

distribution: Weddell Sea (Ringuelet, 1957); South Atlantic (Ringuelet, 1955); North
Atlantic (Dons, 1914); Arctic Ocean (Hadzi, 1951); widespread but in low abundance,
possibly cosmopolitan.

Parafolliculina patagonica (Ringuelet) Ringuelet (1957: 217, fig. 1)

BASIONYM: Claustrofolliculina patagonica Ringuelet (1953b: 243, figs 20-33)

ADDITIONAL REFERENCE: Ringuelet ( 1955: 187)

Fig. 14.8

DESCRIPTION: Lorica c. 140-185 * 65-90 um, slightly irregularly flask-shaped, indistinctly

violet or colourless;ampulla lying, separated from the neck by a bipartite closing membrane;
neck cylindrical, rising at c. 45° from substratum; opening everted. Trophont yellowish,
attached to lorica by a short stalk. Macronucleus 1. Epizoic on bryozoans, isopods and
molluscs.

distribution: Bellingshausen Sea, Weddell Sea, South Atlantic (Ringuelet, 1957).

Parafolliculina violacea (Giard) Dons (1914: 61, 70, pi. 2, fig. 7)

BASIONYM: Folliculina violacea Giard (1888: 315, pi. 20, fig. 6)

SYNONYM: Folliculina telesto Laackmann (1910b: 83, pi. 13, figs 6-9)

ADDITIONAL REFERENCES: Dons (1911: 85), Faure-Fremiet (1936: 1161), Andrews (1942: 91),
Hadzi (1951: 188), Ringuelet (1955: 182; 1957: 215), Mulisch el al. (1986: 5)

Fig. 14.9a, b

DESCRIPTION: Lorica 175-335 (-440) * 50-95 um, bottle-shaped, almost circular in cross
neck straight, often bent in entoecic specimens, on
section, standing upright, pale blue-green;
broad side with a swelling (chamber) that contains a bipartite closing membrane, on narrow
side with a transverse constriction (furrow); opening everted. Trophont to 500 um long.

352
 CILIATES

Figures 14.12-14. Condylostoma and Uronychia species. 14.12, C. granulosum, ventral

sides from life (a) and after protargol impregnation (b) (from Petz et ai, 1995). 14.13, C.
remanei, ventral side from life (a, from Kahl, 1932) and ventral (b) and dorsal (c) sides after
protargol impregnation (from Song & Wilbert, 2002). 14.14, U. transfuga, ventral side from
life (a) and ventral (b) and dorsal (c) sides after protargol impregnation (from Petz et al.,

1995). Scale bars: 12a = 200 urn; 12b = 150 urn; 13 = 100 urn; 14a = 50 um; 14b, c = 30
|xm. Abbreviations: aAM, anterior adoral membranelles; aM, apical membranelles; AZM,
adoral zone of membranelles; CC, caudal cirri; cV, contractile vacuole; Dk, dorsal kineties;
FVC, frontoventral cirri; LMC, left-marginal cirri; Mi, micronucleus; Mn, macronucleus;
pAM, posterior adoral membranelles; TC, transverse cirri; uM, undulating membrane.

353
 CILIATES

16a

Figures 14.15, 16. Diophrys species. 14.15, D. appendiculata, ventral side from life (a, from
Song & Packroff, 1997) and ventral (b) and dorsal (c) sides after protargol impregnation
(from Petz et al., 1995). 14.16, D. oligothrix, ventral side from life (a) and ventral (b) and
dorsal (c) sides after protargol impregnation (from Song & Wilbert, 2002). Scale bars: 15a =
20 pm; 15b, c = 50 urn; 16 = 40 pm. Abbreviations: AZM, adoral zone of membranelles;
CC, caudal cirri; DK, dorsal kineties; FVC, frontoventral cirri; LMC, left-marginal cirri;
Mn, macronucleus (outline); PTC, pretransverse cirri; TC, transverse cirri.

354
 CILIATES

plum-red or blue. Somatic kineties c. 50 longitudinal. Peristomial wings c. 200 urn long,
narrow, equal in size and shape. Macronucleus 1. Epibiotic on algae, bryozoans, isopods and
molluscs.

distribution: Bellingshausen Sea (Ringuelet, 1957); cosmopolitan.

Pseudofolliculina melitta (Laackmann) Dons (1914: 61)

BASIONYM: Folliculina melitta Laackmann (1910b: 80, figs 1-4, 11-15)

ADDITIONAL REFERENCES: Kahl (1932: 473), Hadzi (1951: 156)

Fig. 14.10

description: Lorica 620-770 x 115-145 um, standing upright, usually cylindrical,

sometimes slightly curved, colourless; base occupying posterior third of lorica, with a central
tube; neck containing a closing membrane; opening slightly everted. Trophont posteriorly
with a long stalk. Macronucleus 1; micronuclei 2 or 3. Epizoic on hydroids.

distribution: Indian sector of the Southern Ocean (Laackmann, 1910b).

The population of Folliculina melitta described by Dons (1911) from northern Norway
probably belongs to Pedifolliculina arctica (Hadzi, 1951).

Family Stentoridae

Heterostentor coeruleus Song & Wilbert (2002b: 49, figs 10, 41-44)

Fig. 14.11a, b

description: In vivo c. 200-300 x 60-90 um. Body elongate-cylindrical to broadly fusiform;

apical end slightly head-like; dorsoventrally somewhat flattened. Dark blue pigments
between kineties render cells bright blue. Somatic kineties 100-120, forming a suture on
ventral side. Adoral membranelles c. 50-70, around apical end in an open ©"shape.
Macronucleus ellipsoidal. Movement: slowly gliding.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002b).

Family Condylostomatidae

Condylostoma granulosum Bullington (1940: 191, fig. 7)

ADDITIONAL REFERENCE: Petz et al. (1995: 103)

Figs 14.12a, b; 14.172

description: In vivo c. 65-200 um. Body elongate-ellipsoidal, contractile, rarely

140-590 x

slightly twisted; oral cavity c. 40%

of body length. Somatic kineties 28-65. Adoral
membranelles 123-210, 1-3 short apical membranelles ('cirri') on anterior right of oral
cavity. Colourless cortical granules in interkinetal rows. Macronucleus beaded, composed of
6-13 nodules. Contractile vacuole subterminal (or lacking?).

distribution: Ross Sea, in the region 65.28°-74.00°S and 135.00°-165.05°W, in sea-ice (Petz,
unpublished data); Weddell Sea (Petz et al, 1995); Atlantic (Bullington, 1940).

Condylostoma remanei Spiegel, in Kahl (1932: 456, fig. 75.4, 5)

BASIONYM: Condylostoma caudatum Spiegel (1926: 187, fig. C), due to homonymy with Condylostoma
caudatum Lauterborn (1908)
See Kahl (1932) for synonymy.

ADDITIONAL REFERENCES: Kahl (1928b: 196), Villeneuve-Brachon (1940: 70), Fjeld (1955: 50),
Dragesco (1960: 293), Song & Wilbert (2002b: 52)

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 CILIATES

Fig. 14.13a-c

DESCRIPTION: In vivo 200-1500 * 40-130 [im. Body slender, posteriorly tapering tail-like
(length of tail variable), distinctly flexible, contractile; oral cavity to c. 20% of body length.
About 20-40 somatic kineties forming a suture on posterior ventral side. About 85-200
adoral membranelles; 2-several short apical membranelles ('cirri'). Macronucleus beaded,
composed of 3-19 nodules; micronuclei c. 5. Movement: burrowing.
distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002b); Atlantic,
Mediterranean Sea (Dragesco, 1960); Baltic Sea (Kahl, 1932).

CLASS SPIROTRICHEA
Order Euplotida

Family Uronychiidae

Diophrys appendiculata (Ehrenberg) Schewiakoff (1893: 67)

BASIONYM: Stylonychia appendiculata Ehrenberg (1838: 373, pi. 42, fig. 3)

SYNONYMS: Diophrys marina Dujardin (1841 445, : pi. 10, fig. 4)

Styloplotes appendiculatus (Ehrenberg) Stein (1859b: 62)

Diophrys sp. (Thompson, 1972: 283, fig. 23)

See Curds & Wu (1983) for additional synonyms.

ADDITIONAL REFERENCES: Kahl (1932: 625), Borror (1963: 514), Curds & Wu (1983: 203), Hill
&Borror(1992: 148), Petz et al. (1995: 182), Song & Packroff( 1997: 352)

Figs 14.15a-c; 14.169

description: In vivo 30-120 * 30-70 \im. Body ellipsoidal, slightly indented in region of
caudal cirri, sometimes with a spur-like edge on anterior right, dorsoventrally flattened.
Cirri: 5 or 6 frontoventral, 2 or 3 pretransverse, 4 or 5 transverse, 1-3 left-marginal, 3 caudal
and prominent. Dorsal kineties 5 or 6. Adoral zone composed of 30-37 membranelles that do
not extend onto the right ventral body side. Macronuclear nodules 2 (-5); micronuclei 2—4.
Contractile vacuole 1. Movement: moderately rapid.

distribution: Ross Sea, at 74°02'S 140°01'W, in sea-ice (Petz, unpublished data); Bellings-
hausen Sea (Thompson, 1972); Weddell Sea (Petz et al., 1995); cosmopolitan (Dragesco &
Dragesco-Kerneis, 1986).

Diophrys oligothrix Borror (1965: 559, figs 17-19)

ADDITIONAL references: Czapik(1981: 367), Curds & Wu(1983: 210), Song & Packroff (1997:
351), Song & Wilbert (2002: 54)

Fig. 14.16a-c

description: In vivo 50-106 * 30-60 urn. Body ellipsoidal, without prominent ventral or
dorsal sculpturing, dorsoventrally flattened. Cirri: 5 frontoventral, 2 pretransverse, 5 thick
transverse, 2 or 3 left-marginal, 3 caudal and thick. Dorsal kineties 4 or 5. Adoral zone
composed of 26-45 membranelles, also extending a short distance onto right ventral body
side. Corticalgranules around base of ciliary organelles. Macronuclear nodules 2;
micronuclei 3-6. Movement: crawling rather slowly.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002); Atlantic
(Borror, 1965); Baltic Sea (Czapik, 1981); Pacific (Song & Packroff, 1997).

Diophrys scutum (Dujardin) Kahl (1932: 624, fig. 122.1)

BASIONYM: Ploesconia scutum Dujardin (1841: 437, pi. 10, fig. la; pro parte)

356
 CILIATES

See Curds & Wu (1983) for synonymy.

ADDITIONAL REFERENCES: Curds & Wu (1983: 203), Song & Packroff (1997: 348), Song &
Wilbert (2002: 55)

Fig. 14.17a-c

description: In vivo c. 140-200 * 70-100 urn. Body ellipsoidal, indented posteriorly on right,

Figures 14.17, 18. Diophrys and Aspidisca species. 14.17, D. scutum, ventral side from life
(a) and ventral (b) and dorsal (c) sides after protargol impregnation (from Song & Packroff,
1997). 14.18, A. antarctica, ventral side from life (a, from Petz et al., 1995) and ventral (b)
and dorsal (c) sides after protargol impregnation (from Corliss & Snyder, 1986). Scale bars:
17 = 50 |im; 18 = 20 um. Abbreviations: AZM, adoral zone of membranelles; Fm, frontal
membranelles; FVC, frontoventral cirri; Mn, macronucleus; Ps, prostomial spur; TC,
transverse cirri.

357
 CILIATES

with 2 ventral ridges, dorsoventrally flattened. Cirri: 5-7 frontoventral, 2 pretransverse, 5 or 6

thick transverse, 2 left-marginal, 3 or 4 caudal and thick. Dorsal kineties 5-9. Adoral zone
composed of c. 59-78 membranelles, extending also along anterior third to half of right body

margin. Macronuclear nodules 2 (-7); micronuclei 2-6. Movement: crawling rather quickly.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002); Atlantic
(Borror, 1972); Mediterranean Sea (Dujardin, 1841); Pacific (Song & Packroff, 1997).

Uronychia transfuga (Miiller) Stein (1859a: 129, pi. 4, figs 1-5)

BASIONYM: Trichoda transfuga Miiller (1776: 7, pi. 9, fig. 4)

SYNONYMS: Uronychia antarctica Valbonesi & Luporini (1990a: 365, figs 1-6)

Figures 14.19, 20. Aspidisea species. 14.19, A. crenata, ventral side from life (a) and ventral
(b) and dorsal (c) sides after protargol impregnation (from Song & Wilbert, 2002). 14.20, A.
polypoda, ventral and dorsal sides from life (a, b) and after protargol impregnation (c, d)
(from Song & Wilbert, 2002). Scale bars: 19, 20a, b = 20 urn: 20c, d = 10 urn. Abbreviation:
cV, contractile vacuole.

358
 CILIATES

Uronychia binucleata Young (1922: 360, figs 4-6)

Uronychia uncinata Taylor (1928: 3, fig. 1)

ADDITIONAL REFERENCES: Kent (1882: 797), Kahl (1932: 627), Borror (1963: 128), Curds & Wu
(1983:220), Hill (1990: 100), Petz et al. (1995: 173)

Figs 14.14a-c; 14.162

DESCRIPTION: In vivo usually 80-110 * 50-65 um. Body ±rectangular, indented in area of
transverse and left-marginal cirri, with spur-like protrusions anteriorly and posteriorly,

dorsoventrally flattened. Cirri very prominent: 6 frontoventral, 4 or 5 transverse, 3 left-

marginal, 3 caudal. Dorsal kineties 6. Adoral zone of membranelles bipartite, with
1 1 anterior and 5 posterior membranelles. Undulating membrane almost surrounding a very

large oral area. Macronuclear nodules 2, elongate-ellipsoidal; micronucleus 1. Contractile

vacuole 1. Movement: usually jumping and swimming very fast, sometimes slowly and
wobbly, resting in-between.

DISTRIBUTION: Ross Sea, in and 164.10°E-134.85°W, in sandy

the region 66.00°-74.70°S
sediments, seawater and sea-ice (Valbonesi &
1990a; Petz, unpublished data);
Luporini,
Atlantic sector of Southern Ocean (Song & Wilbert, 2002b); Weddell Sea (Fenchel & Lee,
1972; Petz et al., 1995); Greenland Sea (Agatha et al., 1993); widespread and cosmopolitan.

Specimens with a nodulated, C-shaped or horseshoe-shaped macronucleus belong to U.

heinrothi Buddenbrock (see Petz et al., 1995).

Family Aspidiscidae

Aspidisca antarctica Corliss & Snyder (1986: 44, fig. 7)

ADDITIONAL REFERENCE: Petz et al. (1995: 172)

Fig. 14.18a-c

description: In vivo 45-55 x 30-40 um. Body D-shaped, widest at or behind mid-body,
posteriorly on left with a distinct thorn-like projection (prostomial spur) associated with
indentation of cell, sometimes right border of indentation also spur-like, with 4 conspicuous
dorsal ridges, dorsoventrally flattened. Cirri: 7 or 8 frontoventral, 5 or 6 transverse. Dorsal
kineties 4, each with 4-10 dikinetids. Frontal membranelles 3; adoral membranelles c. 9-12.
Macronucleus horseshoe-shaped; micronucleus 1. Contractile vacuole 1. Movement: slowly
crawling, rotating about main body axis when swimming.

distribution: Ross Sea, in the region 66.00°-74.03°S and 134.85°-165.02°W, in sea-ice (Petz,
unpublished data); Bellingshausen Sea (Kloser, 1990); Weddell Sea (Corliss & Snyder, 1986;
Petz et al, 1995).

Aspidisca crenata Fabre-Domergue (1885: 561, pi. 28, fig. 7)

ADDITIONAL REFERENCES: Kahl (1932: 649), Song & Wilbert (2002: 58)

Fig. 14.19a-c

DESCRIPTION: In vivo 50-70 x 40-55 um. Body broadly D-shaped, a triangular thorn-like
projection covering adoral membranelles (Song & Wilbert, 2002); other populations with
distinct spur-like protrusion on left-lateral margin (Fabre-Domergue, 1885; Kahl, 1932),
dorsoventrally flattened, with 3 or 4 longitudinal dorsal ridges. Cirri: 7 frontoventral, 5-7
transverse. Dorsal kineties 4, densely ciliated (c. 20 dikinetids each). Frontal membranelles
4; adoral membranelles 17-19. Macronucleus C-shaped. Contractile vacuole 1. Movement:
crawling.

DISTRIBUTION: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002); Atlantic (Kahl,
1932).

359
 CILIATES

Aspidisca polypoda (Dujardin) Kahl (1932: 646, fig. 125.30, 31)

BASIONYM: Coccudina polypoda Dujardin (1841: 447, pi. 10, fig. 3)

ADDITIONAL REFERENCES: Wu & Curds (1979: 19), Song & Wilbert (2002: 57)

Fig. 14.20a-d

DESCRIPTION: In vivo 25-55 x 20-45 um. Body triangular to D-shaped, without a thorn-like
projection, dorsally distinctly vaulted and with 7 or 8 prominent longitudinal ridges,
dorsoventrally flattened. Cirri: 7 frontoventral, 5 or 6 transverse. Dorsal kineties 4, each with
4-7 dikinetids. Frontal membranelles 3 or 4; adoral membranelles 8-15. Macronucleus
horseshoe-shaped; micronucleus 1. Contractile vacuole 1. Movement: crawling moderately
quickly.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002); North Sea
(Kahl, 1932); Mediterranean Sea (Dujardin, 1841).

Aspidisca quadrilineata Kahl (1932: 647)

ADDITIONAL REFERENCE: Song & Wilbert (2002: 58)

Fig. 14.21a-c

description: In vivo 30-40 * 25-35 urn. Body broadly D-shaped, without a thorn-like
projection, with 4 indistinct dorsal ridges, dorsoventrally flattened. Cirri: 7 frontoventral, 5-7
7-16 dikinetids. Frontal membranelles 4 or 5; adoral
transverse. Dorsal kineties 4, each with
membranelles 12-14. Macronucleus horseshoe-shaped. Contractile vacuole 1. Movement:
crawling slowly to moderately quickly.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002); North Sea,
Baltic Sea (Kahl, 1932).

Aspidisca terranovae Valbonesi (1996: 378, figs 1-6)

Fig. 14.22a, b

description: Fixed specimens 30-40 x 20-30 um. Body ellipsoidal, dorsoventrally

flattened, with a distinct thorn-like projection subequatorially on left-lateral margin; dorsal
surface smooth. Cirri: 7 frontoventral, 5 transverse. Dorsal kineties 4 (or 5), each with 4-7
dikinetids. Frontal membranelles 4; adoral membranelles 13 or 14. Macronucleus horseshoe-
shaped; micronucleus 1. Thigmotactic, such that crawling specimens cannot be easily
removed with a pipette.
distribution: Ross Sea (Valbonesi, 1996).

Family Gastrocirrhidae

Cytharoides balechi Tuffrau (1974: 313, figs 1-8)

SYNONYM: Cytharoides australis Agatha, Wilbert, Spindler & ElbrShter (1990: 224, fig. 3)

ADDITIONAL REFERENCES: Curds & Wu (1983: 244), Agatha et al. (1990: 223), Petz et al. (1995:
178), Song & Wilbert (2000a: 221)

Figs 14.23a-d; 14.170

DESCRIPTION: In vivo 120-220 x 90-120 um. Body elongate, anteriorly ±obliquely truncate,
posteriorly asymmetrically tapering, dorsoventrally flattened; buccal field large, with many
indistinct dorsal ridges. Cirri: 10 frontoventral, 5 transverse, 2 left-marginal, 8-16 caudal.
Dorsal kineties 13-38. Adoral membranelles 61-144. Macronucleus C-shaped to somewhat
3-shaped; micronucleus 1. Contractile vacuole 1. Dorsal silverline system of double-patella
type. Movement: slowly crawling, usually in tight circles.

360
 CILIATES

distribution: Ross Sea (Coppellotti et al, 1992; Gowing et al, 2001); Bellingshausen Sea
(Tuffrau, 1974); Weddell Sea (Agatha et al, 1990; Petz et al, 1995; Song & Wilbert, 2000a).

Family Euplotidae

Euplotes acanthodus Petz, Song & Wilbert (1995: 183, fig. 55)

Fig. 14.24a-c

description: In vivo 90-130 * 60-85 um. Body triangular to broadly obovoid, with a
prominent spur on anterior right and c. 8 indistinct dorsal ridges, dorsoventrally flattened.
Cirri: 10 frontoventral, 5 transverse, 2 left-marginal, 2 caudal. Dorsal kineties 12 (15-20
dikinetids in mid-dorsal rows). Adoral membranelles 60-76. Macronucleus C-shaped.
Contractile vacuole 1. Dorsal silverline system of double-eurystomus type. Movement:
crawling slowly on substratum.

21a

22a

Figures 14.21, 22. Aspidisca species. 14.21, A. quadrilineata, ventral side from life (a) and
ventral (b) and dorsal (c) sides after protargol impregnation (from Song & Wilbert, 2002).
14.22, A. terranovae, ventral (a) and dorsal (b) sides after silver impregnation (from
Valbonesi, 1996). Scale bars: 21 = 10 um; 22 = 20 um. Abbreviation: SI, silverline.

361
 CILIATES

distribution: Ross Sea, in the region 65.95°-74.00°S and 135.00°-150.08°W, in sea-ice (Petz,
unpublished data); Weddell Sea (Petz et al, 1995).

Euplotes antarcticus Fenchel & Lee (1972: 234, fig. 3)

ADDITIONAL REFERENCE: Petz et al. (1995: 186)

Figs 14.25a-c; 14.166

Figures 14.23, 24. Cytharoides and Euplotes species. 14.23, C. balechi, ventral side from life
(a), ventral (b) and dorsal (c) sides after protargol impregnation and detail of dorsal silverline
system (d) after silver nitrate impregnation (from Petz et al., 1995). 14.24, E. acanthodus,
ventral side from life (a) and after protargol impregnation (b), dorsal side with silverline system
(c) after silver nitrate impregnation (from Petz et al., 1995). Scale bars: 23a-c = 50 urn; 24 = 30
urn. Abbreviations: Bf, buccal field; CC. caudal cirri; S, spur; uM. undulating membrane.

362
 CILIATES

Figures 14.25, 26. Euplotes species. 14.25, E. antarcticus, ventral side from life (a) and
after protargol impregnation (b), dorsal side with silverline system (c) after silver nitrate
impregnation (from Petz et ah, 1995). 14.26, E. balteatus, ventral side from life (a), ventral
(b) and dorsal (c) sides after protargol impregnation and detail of dorsal silverline system (d)
after silver nitrate impregnation (from Song & Wilbert, 2002). Scale bars: 25 = 40 |im; 26a-
c = 30 urn.

363
 CILIATES

Figures 14.27-29a. Euplotes species. 14.27, E. euryhalinus, ventral side from life (a) and
ventral (b) and dorsal (c) sides after protargol impregnation. 14.28, E. focardii, ventral side
after silver carbonate impregnation (from Serrano et al., 1992). 14.29a, E. rariseta, ventral
side from life (from Petz et al, 1995). Scale bars: 27, 28 = 30 urn; 29a = 20 ^im.
Abbreviations: F, fibres; Mi, micronucleus.

364
 CILIATES

description: In vivo 90-145 x 30-80 urn. Body elongate-ellipsoidal; anterior ventral surface
distinctly protruding, with 3 or 4 inconspicuous ventral and 6-10 low dorsal ridges,
dorsoventrally flattened. Cirri: 10 frontoventral, 5 transverse, 2 left-marginal, 3-6 caudal.
Dorsal kineties 8-14 (11-21 dikinetids in mid-dorsal rows). Adoral membranelles 30-70.
Macronucleus question-mark-shaped; micronucleus 1. Contractile vacuole 1. Dorsal
silverline system of double-eurystomus type. Movement: crawling very slowly on
substratum, remaining stationary for long periods.

distribution: Ross Sea, in the region 71.03°-74.03°S and 1 35.00°- 150.40°W, in sea-ice (Petz,
unpublished data); Weddell Sea (Fenchel & Lee, 1972; Corliss & Snyder, 1986; Petz et al,
1995).

Euplotes balteatus (Dujardin) Kahl (1932: 635, fig. 123.7, 8)

BASIONYM: Ploesconia balteata Dujardin (1841: 437, pi. 10, fig. 12)

ADDITIONAL REFERENCES: Curds (1975: 22), Tuffrau (1964: 143), Song & Wilbert (2002: 55)

Fig. 14.26a-d

DESCRIPTION: In vivo 30-150 * 30-105 urn. Body broadly ellipsoidal, with 3 or 4

conspicuous ventral ridges and 6 or 7 dorsal ridges, dorsoventrally flattened. Cirri: 10
frontoventral, 5 transverse, 1 or 2 left-marginal, 2 caudal. Dorsal kineties 8-10 (c. 9-14
dikinetids in mid-dorsal rows). Adoral membranelles 25-80. Macronucleus C- to horseshoe-
shaped; micronucleus 1. Contractile vacuole 1. Dorsal silverline system of double-
eurystomus type (sometimes slightly irregular). Movement: crawling slowly to moderately
quickly.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002); North Sea,
Baltic Sea, Mediterranean Sea (Kahl, 1932); also endozoic in sea urchins (Berger, 1965).

Euplotes euryhalinus Valbonesi & Luporini (1990b: 50, figs 8-12)

Figs 14.27a-c; 14.167

description: In vivo 60-100 * 40-60 urn.Body broadly ellipsoidal, with a small spur-like
projection on anterior right in c. 50% of specimens, 1 or 2 conspicuous ventral and 6-9
dorsal ridges, dorsoventrally flattened; pellicle rigid. Cirri: 10 frontoventral, 5 or 6
transverse, 2 or 3 left-marginal, 2 or 3 caudal. Dorsal kineties 1 1 (or 12) (13-21 dikinetids in
mid-dorsal rows). Adoral membranelles 26-35. Macronucleus horseshoe- to C-shaped, rarely
inverted J-like; micronucleus 1. Contractile vacuole 1. Dorsal silverline system of double-
eurystomus type. Movement: crawling moderately quickly, interrupted by resting. Specimens
can encyst rapidly.

distribution: Brackish pools on shore of Ross Sea (Valbonesi & Luporini, 1990b).

This species has also been found in freshwater lakes in Victoria Land, in the region 74.33°-
74.70°S and 164.12°-165.12°E (Petz & Valbonesi, unpublished data), and on Byers Peninsula,
Livingston I. (South Shetland Is.) at about 62°38'S 61°04'W (Petz & (jfesada, unpublished data).

Euplotes focardii Valbonesi & Luporini (1990c: 58, figs 1-12)

additional REFERENCES: Serrano et al. (1992: 2088), Valbonesi & Luporini (1993: 489)

Fig. 14.28

description: Fixed specimens 38-110 * 30-92 um. Body ellipsoidal to roundish, dorsal
surface smooth except for 2 ridges near left margin (undernourished cells with c. 6 additional
inconspicuous ridges), dorsoventrally flattened. Cirri: 10 frontoventral, 5 transverse, 2 left-
marginal, 2 caudal. Dorsal kineties 9-12 (c. 13-22 dikinetids in mid-dorsal rows). Adoral
membranelles 39-72. Macronucleus horseshoe-shaped; micronucleus 1. Contractile vacuole
1. Dorsal silverline system slightly irregular and of double-eurystomus to indistinct double-

patella type. Movement: crawling.

365
 CILIATES

distribution: Ross Sea (Valbonesi & Luporini, 1990c, 1993).

Euplotes rariseta Curds, West & Dorahy (1974: 97, figs 2-4)

SYNONYMS: Euplotes algivorus Agatha, Wilbert, Spindler & Elbrahter (1990: 226, fig. 4)
Euplotes nobilii Valbonesi & Luporini (1990b: 48, figs 3-5)
Euplotes sp. (Thompson, 1972: 282, fig. 21)

additional REFERENCES: Curds (1975: 42), Wilbert & Kahan (1981: 88), Valbonesi & Luporini
(1990b: 51), Petz et al (1995: 191), Song & Packroff( 1997: 346)

Figs 14.29a-d; 14.171

DESCRIPTION: In vivo 30-67 * 25^45 um. Body ellipsoidal to slightly triangular, with a spur-
like projection on anterior right, posterior margin usually transparent, with c. 3
inconspicuous ventral and c. 6 ±prominent dorsal ridges, dorsoventrally flattened. Cirri: 10
frontoventral, 5 transverse, 1 (sometimes 2 very closely set) left-marginal, 2 caudal. Dorsal
kineties 5-8 (c. 6-14 dikinetids in mid-dorsal rows). Adoral membranelles 18-37.
Macronucleus C- to S-shaped; micronucleus 1. Contractile vacuole 1. Dorsal silverline
system of double-patella type. Movement: crawling moderately quickly, jerking slightly.

distribution: Ross Sea (Valbonesi & Luporini, 1990b; Albergoni et al, 2000); Bellings-
hausen Sea (Thompson, 1972); Weddell Sea (Agatha et al, 1990; Petz et al, 1995); Atlantic
(Curds et al, 1974); Indian Ocean (Dallai et al, 1987); Red Sea (Wilbert & Kahan, 1981);
Pacific (Song & Packroff, 1997).

Euplotes sigmolateralis Agatha, Spindler & Wilbert (1993: 263, fig. 3)

Fig. 14.30a, b

description: Fixed specimens 78-149 x 46-113 um. Body elongate; left margin distinctly
convex; right margin sigmoidal; dorsoventrally flattened. Cirri: 7 frontoventral (cirrus 112
massive), 5 transverse, left-marginal, 2 caudal. Dorsal kineties 7-13 (15-23 dikinetids in
mid-dorsal rows). Adoral membranelles 47-67. Macronucleus sausage-shaped.

distribution: Weddell Sea, Greenland Sea (Agatha et al, 1993).

Order Tintinnida

Family Codonellidae

Tintinnopsis bacillaria Hada (1970: 30, fig. 46)

additional REFERENCES: Sassi & Melo (1986: 65), Song & Xu (1999: 35)

Fig. 14.31

description: Lorica 63-95 * 30-46 um, bullet-shaped, with agglutinated diatoms and other
foreign particles, sometimes hyaline and without particles.

distribution: Atlantic sector of the Southern Ocean (Sassi & Melo, 1986; Wasik &
Mikolajczyk, 1994a); Weddell Sea (Song & Xu, 1999); South Atlantic (Hada, 1970).

Tintinnopsis buetschlii Daday (1887: 556, pi. 20, figs 4, 5)

SYNONYMS: Tintinnopsis major Meunier (1910: 138, pi. 12, fig. 1)

Tintinnopsis loricata (Brandt) Kofoid & Campbell (1929: 39, fig. 60)

additional REFERENCES: Brandt (1907: 145), Balech (1948: 10)

Fig. 14.32

366
 CILIATES

description: Lorica 87-170 * 69-110 um, elongate-campanulate; anterior half of lorica

annulated, apically everted funnel-like, antapically tapering (sometimes rounded); wall with
agglutinated mineral particles. Trophont with c. 20 adoral membranelles. Macronuclear
nodules 2. Contractile vacuole 1.

distribution: Atlantic sector of the Southern Ocean (Wasik & Mikolajczyk, 1990); Atlantic
(Meunier, 1910; Balech, 1948); Mediterranean Sea (Daday, 1887); Java Sea (Brandt, 1907);
Pacific (Yoo & Kim, 1990).

Tintinnopsis lobiancoi Daday (1887: 553, pi. 19, fig. 27)

SYNONYM: Tintinnopsis lata Meunier (1910: 141, pi. 12, figs 19-22)

29b

T^fi^&

30a

Figures 14.29b-32. Euplotes and Tintinnopsis species. 14.29b-d, E. rariseta, ventral (b) and
dorsal (c) sides after protargol impregnation, and dorsal silverline system (d, as E. nobilii)
impregnation (b, c, from Petz et al, 1995; d, from Valbonesi & Luporini,
after silver nitrate
1990b). 14.30, E. sigmolateralis, ventral (a) and dorsal (b) sides after protargol impregnation
(from Agatha et al., 1993). 14.31, T. bacillaria, lorica (from Hada, 1970). 14.32, T.

buetschlii, lorica with contracted trophont (from Meunier, 1910). Scale bars: 29, 31 = 20 ^m;
30, 32 = 30 um. Abbreviations: Mn, macronuclear nodules; II/2, cirrus 112.

367
 CILIATES

Figures 14.33-40. Tintinnopsis, Stenosemella and Codonellopsis species. 14.33, T. lobiancoi,

lorica (from Kofoid & Campbell, 1929). 14.34, T. parva, lorica shapes (a, from Balech, 1948;
b, as T. glans, from Meunier, 1919). 14.35, T. petasa, lorica (from Hada, 1970). 14.36,
S. avellana, lorica (from Meunier, 1919). 14.37, S. oliva, lorica (from Meunier, 1910). 14.38,

S. perpusilla, lorica (from Hada, 1970). 14.39, C. balechi, lorica (as C. antarctica, from

Balech, 1971). 14.40, C. gauss i, typical (a, from Laackmann, 1907) and coxlielliform loricae
(b, from Alder, 1999). Scale bars: 33 = 40 urn; 34-37 = 10 jam; 38 = 5 urn; 39 = 20 urn; 40 =
50 um. Abbreviations: B, bowl; C, collar; L, lid.

368
 CILIATES

ADDITIONAL REFERENCES: Brandt (1907: 160), Meunier (1910: 138), Kofoid & Campbell (1929:
38), Alder (1999: 344)

Fig. 14.33

description: Lorica 104-390 * 40-80 um, tubular, antapically broadly rounded; wall with
agglutinated mineral particles. Trophont with c. 20 adoral membranelles. Macronuclear
nodules 2. Contractile vacuole 1.

distribution: Atlantic sector of the Southern Ocean (Wasik & Mikolajczyk, 1990, 1994a);
South Atlantic (Alder, 1999); Indian Ocean, Atlantic, North Sea (Brandt, 1907); Kara Sea
(Meunier, 1910); Baltic Sea (Wasik, 1998); Mediterranean Sea (Daday, 1887).

Tintinnopsis parva Merkle (1909: 150, pi. 2, figs 9-1 1)

SYNONYM: Tintinnopsis glans Meunier (1919: 29, pi. 22, figs 35, 36)

See Alder (1999) for additional synonyms.

ADDITIONAL REFERENCES: Balech(1948: 13), Hada(1970: 29), Alder (1999: 344)

Fig. 14.34a, b

description: Lorica 28-42 x 22-28 um, obconical; apical end slightly to distinctly
narrowed; collar indistinct (covered in particles) or lacking; posteriorly usually pointed; wall
with agglutinated mineral particles. Trophont with 2 macronuclear nodules; micronuclei 2.

distribution: Indian sector of the Southern Ocean (Hada, 1970); South Atlantic (Balech,
1948); North Sea (Meunier, 1919); Baltic Sea (Merkle, 1909).

One specimen was reported by Hada (1970) to be antapically broadly rounded.

Tintinnopsis petasa Hada (1970: 30, fig. 45)

Fig. 14.35

description: Lorica c. 30 * 30 um, obconical, apically everted funnel-like, antapically

narrowly rounded; wall thin, with sparsely agglutinated mineral particles.

distribution: Probably in the Indian sector of the Southern Ocean, but exact location not
provided (Hada, 1970).

Family Codonellopsidae

Codonellopsis balechi Hada (1970: 31)

BASIONYM: Codonellopsis frigida Balech (1958a: 78, pi. 1, figs 4-6), due to homonymy with Codonellopsis
Hada (1932)
frigida

SYNONYM: Codonellopsis antarctica Balech (1971: 168, figs 714, 715)

additional reference: Barria de Cao (1987: 276)

Fig. 14.39

description: Lorica 69-136 * 45-80 p.m; collar cylindrical, often short, c. 28-50 um wide,
annulated, hyaline; bowl usually obovoid (sometimes antapically broadly rounded), with
agglutinated mineral particles and broken diatom frustules.

distribution: Bellingshausen Sea (e.g. Balech, 1973a); Atlantic sector of the Southern
Ocean (e.g. Wasik & Mikolajczyk, 1990); Weddell Sea (Barria de Cao, 1987; Alder &
Boltovskoy, 1991b); South Atlantic (Alder, 1999).

369
 CILIATES

Codonellopsis gaussi (Laackmann) Kofoid & Campbell (1929: 79, fig. 164)

BASIONYM: Codonella gaussi Laackmann (1907: 239, fig. 12)

SYNONYM: Leprotintinmts gaussi (Laackmann) Laackmann (1910a: 407, pi. 47, figs 1-4)

ADDITIONAL REFERENCES: Balech (1947: 85; 1973a: 28), Sassi & Melo (1986: 67), Alder (1999:
345)

Figs 14.40a, b; 14.189

description: Lorica 80-180 * 35-60 um; collar cylindrical, variable in length, annulated,
hyaline; bowl distinctly dilated in the middle, terminating in an antapical horn (length
variable), with agglutinated foreign particles (often diatoms). Coxlielliform lorica annulated,
hyaline.

distribution: Indian sector of the Southern Ocean (Laackmann, 1907; Hada, 1970; Grey et
al, 1997); Pacific sector of the Southern Ocean (Balech, 1958b; Tumantseva, 1989); Ross
Sea (e.g. Monti & Fonda Umani, 1995; Fonda Umani et al, 1998); Bellingshausen Sea
(Balech, 1947, 1973a; Alder & Boltovskoy, 1991a); Atlantic sector of the Southern Ocean
(e.g. Wasik & Mikolajczyk, 1990; Thompson et al, 1999); Weddell Sea (e.g. Balech & El-
Sayed, 1965; Boltovskoy & Alder, 1992; Petz et al, 1995); South Atlantic (Alder, 1999).

Codonellopsis glacialis (Laackmann) Kofoid & Campbell (1929: 79, fig. 162)
BASIONYM: Codonella glacialis Laackmann (1907: 239, fig. 13)

SYNONYM: Leprotintinnus glacialis (Laackmann) Laackmann (1910a: 408, pi. 47, figs 5-8)

ADDITIONAL REFERENCE: Petz et al. (1995: 144)

Figs 14.41a-e; 14.163

description: Lorica 55-125 * 33-53 urn, bullet-shaped; collar hyaline, annulated; bowl with
agglutinated mineral and biogenic particles, rarely without agglutinated particles. Trophont
with 25-29 somatic kineties (usually 8 shorter and 2 distinctly elongate ciliary rows in
lateral field, 7 in left and 7 in right field, 1 ventral kinety, 1 dorsolateral kinety). Adoral zone
of membranelles (polykinetid zone) composed of 17-18 external and 1 internal adoral
membranelle in a closed spiral around apical end. Macronuclear nodules 4. Contractile
vacuole 1. Movement: slow to moderately fast.

distribution: Indian sector of the Southern Ocean (Laackmann, 1907; Hada, 1970; Grey et
al, 1997); Ross Sea (e.g. Monti & Fonda Umani, 1995; Fonda Umani et al, 1998);
Bellingshausen Sea (Balech, 1958a; Kloser, 1990); Atlantic sector of the Southern Ocean
(e.g. Wasik & Mikolajczyk, 1990); Weddell Sea (e.g. Balech & El-Sayed, 1965; Garrison &
Buck, 1989a; Boltovskoy & Alder, 1992; Petz et al, 1995).

Balech (1973a) and Alder (1999) considered this species to be synonymous with C. gaussi.
However, without information on the infraciliature this is premature.

Codonellopsis morchella (Cleve) Jorgensen (1924: 100, fig. Ill)

BASIONYM: Codonella morchella Cleve (1900a: 969, fig. [2])

SYNONYM: Codonellopsis turgescens Kofoid & Campbell (1929: 90, fig. 155)
See Alder ( 1 999) for additional synonyms.

ADDITIONAL REFERENCES: Brandt (1907: 124), Kofoid & Campbell (1929: 83), Alder (1999: 345)

Fig. 14.42a, b

DESCRIPTION: Lorica 68-140 * 50-70 um; collar cylindrical, annulated, c. 30 urn wide,
frequently with 2 small fenestrae; bowl obovoid to almost globular, posteriorly rounded, with
agglutinated mineral particles. Trophont with 8 macronuclear nodules.

distribution: Indian sector of the Southern Ocean (Laackmann, 1910a); Pacific sector of the
Southern Ocean (Mamaeva, 1986); cosmopolitan.

370
 CILIATES

Figure 14.41. Codonellopsis glacialis. Left side from life (a), left (b) and right (c) sides
after protargol impregnation and typical (d) and ptychocyliform lorica (e) (from Petz et ai,
1995). Scale bars: a-c = 30 ^m; d, e = 15 um. Abbreviations: AZM, adoral zone of
membranelles; Dl, dorsolateral kinety; eAM, external adoral membranelles; iAM, internal
adoral membranelle; L, lateral field of kineties; Lf, left field of kineties; pM, paroral
membrane; Rf, right field of kineties; Vk, ventral kinety.

371
 CILIATES

45a 46a 47

Figures 14.42-51. Codonellopsis, Laackmanniella, Coxliella, Acanthostomella, Epiplocylis,

Epiplocyloides and Ptychocylis species. 14.42, Codonellopsis morchella, lorica shapes (a, from
Cleve, 1900; b, from Brandt, 1906). 14.43, Codonellopsis pusilla, lorica (from Cleve, 1900).
14.44, Codonellopsis soyai, lorica (from Hada, 1970). 14.45, Laackmanniella naviculaefera,
typical (a, from Petz et al., 1995) and coxlielliform loricae (b, as Cyttarocylis minor, from
Laackmann, 1907). 14.46, Coxliella frigida, lorica shapes (a, from Laackmann, 1907; b, from
Laackmann, 1910a). 14.47, Coxliella intermedia, lorica (from Laackmann, 1907). 14.48,
Acanthostomella norvegica, lorica with contracted trophont (from Meunier, 1910). 14.49,
Epiplocylis undella, lorica (from Ostenfeld & Schmidt, 1902). 14.50, Epiplocyloides
antarctica, lorica and outline of apparently fixed trophont (from Balech, 1962b).

372
 CILIATES

Codonellopsis pusilla (Cleve) Jorgensen (1924: 98)

BASIONYM: Codonella pusilla Cleve (1900a: 970, fig. [3])

ADDITIONAL REFERENCES: Balech(1959: 25; 1971: 170), Sassi & Melo (1986: 67)

Figs 14.43; 14.190

description: Lorica 38-57 x 29—40 urn, clavate; collar cylindrical or slightly narrowed
anteriorly, hyaline, annulated, with few fenestrae; bowl subspherical to slightly tapering,
covered with agglutinated particles (often coccoliths, e.g. of Emiliania huxleyi). Trophont
with 2 macronuclear nodules.

distribution: Indian sector of the Southern Ocean (Scott, unpublished data); Atlantic sector
of the Southern Ocean (Balech, 1971; Sassi & Melo, 1986); Atlantic (Cleve, 1900a; Balech,
1971); White Sea (Burkovsky et al., 1974); Mediterranean Sea (Balech, 1959); Pacific
(Balech, 1962a).

Codonellopsis soyai Hada (1970: 31, fig. 48)

Fig. 14.44

description: Lorica 50-52 * 32-38 urn; collar cylindrical, annulated, with many small
fenestrae; bowl obovoid to almost globular; wall usually smooth, sometimes with agglut-
inated coccoliths.

distribution: Indian sector of the Southern Ocean (Hada 1970).

Probably conspecific with Codonellopsis pusilla (see above).

Laackmanniella naviculaefera (Laackmann) Kofoid & Campbell (1929: 91, fig. 182)

BASIONYM: Codonella naviculaefera Laackmann (1907: 239, fig. 10)

SYNONYMS: Cyttarocylis minor Laackmann (1907: 237, fig. 7)

Codonella prolongata Laackmann (1907: 239, fig. 11)

Coxliella minor (Laackmann) Laackmann (1910a: 397, pi. 45, figs 12-15)

Leprotintinnus naviculaeferus (Laackmann) Laackmann (1910a: 402, pi. 46, figs 1-9)

Leprotintinnus prolongatus (Laackmann) Laackmann (1910a: 403, pi. 46, figs 10-12)

Laackmanniella prolongata (Laackmann) Kofoid & Campbell (1929: 91, fig. 183)

ADDITIONAL REFERENCES: Balech (1947: 85; 1958b: 398), Sassi & Melo (1986: 69), Barria de
Cao (1987: 277), Petz etal. (1995: 151), Alder (1999: 357)
Figs 14.45a, b; 14.168a, b

description: Lorica 100-310 x 30-55 urn; collar long, cylindrical, annulated, apically
slightly dilated or straight, hyaline; bowl slightly inflated, posteriorly narrowed, antapically
open, covered with agglutinated mineral particles, some diatoms and detritus. Coxlielliform
lorica annulated, hyaline, without agglutinated particles. Trophont with 4 macronuclear
nodules and 2 micronuclei.

distribution: Indian sector of the Southern Ocean (Laackmann, 1907; Hada, 1970; Grey et
al.,1997); Pacific sector of the Southern Ocean (Balech, 1958b); Ross Sea (e.g. Hopkins,
1987; Monti & Fonda Umani, 1995); Bellingshausen Sea (e.g. Balech 1947, 1973a; Kloser,
1990); Atlantic sector of the Southern Ocean (e.g. Sassi & Melo, 1986; Thompson et al.,
1999); Weddell Sea (e.g. Balech & El-Sayed, 1965; Heinbokel & Coats, 1986; Boltovskoy &
Alder, 1992; Petz etal., 1995); South Atlantic (Alder, 1999).

Continued ... 14.51, Ptychocylis ostenfeldi, lorica (from Kofoid & Campbell, 1929). Scale
bars: 42a, 45b, 47, 51 = 40 urn; 42b, 46b, 49, 50 = 30 urn; 43, 44 = 20 urn; 45a = 25 urn;
46a = 50 p.m; 48 = 10 urn. Abbreviations: F, fenestra; H, horn; Ic, inner collar; Oc, outer
collar; R, rim; Re, reticulum.

373
 CILIATES

Stenosemella a veil ana (Meunier) Kofoid & Campbell (1929: 69, fig. 134)
BASIONYM: Tintinnopsis avellana Meunier (1919: 30, pi. 22, fig. 37)

ADDITIONAL REFERENCES: Balech (1948: 17), Hada(1961: 143), Alder (1999: 345)

Fig. 14.36

description: Lorica 38-68 * 32-68 um, ellipsoidal to subglobular; apical opening 20-24 um
diameter; lid (closing apparatus) sometimes with agglutinated particles; collar short and
sometimes covered with particles; bowl with agglutinated mineral particles.

DISTRIBUTION: Atlantic sector of the Southern Ocean (Souto Jankilevich, 1981); South &
Atlantic (Balech, 1948); North Sea (Meunier, 1919); South Pacific (Hada, 1961).

Stenosemella oliva (Meunier) Kofoid & Campbell (1929: 70, fig. 135)
BASIONYM: Tintinnopsis oliva Meunier (1910: 144, pi. 13, figs 9-13)

Fig. 14.37

DESCRIPTION: Lorica 46-60 * 36-44 um, obovoid; collar short and hyaline; lid and bowl with
agglutinated mineral particles. Trophont with 1 macronucleus.

distribution: Atlantic sector of the Southern Ocean (Wasik & Mikolajczyk, 1994a); North
Atlantic (Meunier, 1910); White Sea (Burkovsky et ai, 1974).

Stenosemella perpusilla Hada (1970: 30, fig. 47)

Fig. 14.38

description: Lorica c. 13 * 11 um; collar short and hyaline; bowl globular, covered with
agglutinated particles.

distribution: Indian sector of the Southern Ocean (Hada, 1970).

Family Metacylididae

Coxliella frigida (Laackmann) Laackmann (1910a: 392, pi. 44, figs 1-8)

BASIONYM: Cyttarocylis frigida (Laackmann 1907: 237, fig. 6)

ADDITIONAL REFERENCES: Kofoid & Campbell (1929: 99), Balech (1947: 86; 1958b: 398), Sassi
&Melo(1986: 71)

Fig. 14.46a, b

description: Lorica 110-345 * 75-110 um, cylindrical, irregularly annulated, apically

sometimes everted, posteriorly broadly rounded; rim serrated. Trophont with 2 macronuclear
nodules and 2 micronuclei.

distribution: Indian sector of the Southern Ocean (Laackmann, 1907); Pacific sector of the
Southern Ocean (Balech, 1958b); Ross Sea (e.g. Monti & Fonda Umani, 1995; Fonda Umani
et ai, 1998); Bellingshausen Sea (Balech, 1947; Barria de Cao, 1987; Kloser, 1990); Weddell
Sea (e.g. Balech & El-Sayed, 1965; Heinbokel & Coats, 1986; Boltovskoy et ai, 1989).

This is listed as a synonym of Cymatocyl is convallaria by Alder (1999).

Coxliella intermedia (Laackmann) Laackmann (1910a: 394, pi. 45, figs 5-1 1)

BASIONYM: Cyttarocylis intermedia Laackmann (1907: 238, fig. 8)

ADDITIONAL REFERENCES: Balech ( 1 958a: 79; 1973a: 29), Sassi & Melo (1986: 71)

Fig. 14.47

374
 CILIATES

description: Lorica 119-205 * 54-84 jim, cylindrical, irregularly annulated, posteriorly

narrowly rounded; wall hyaline, brownish. Trophont with 2 macronuclear nodules and 2
micronuclei.

distribution: Indian sector of the Southern Ocean (Laackmann, 1907); Ross Sea (Petz,
2001); Bellingshausen Sea (Balech, 1958a, 1973a; Kloser, 1990); Atlantic sector of the
Southern Ocean (Sassi & Melo, 1986; Wasik & Mikolajczyk, 1994a); Weddell Sea (Balech
&E1-Sayed, 1965).
Alder (1999) considers this to be conspecific with Cymatocvlis drygalskii (Xystonellidae, see
p. 380).

Family Ascampbelliellidae

Acanthostomella norvegica (Daday) Jorgensen (1927: 13, fig. 23)

BASIONYM: Amphorella norvegica Daday (1887: 543)

See Kofoid & Campbell (1929) for synonymy.

ADDITIONAL REFERENCES: Balech(1971: 174), Alder (1999: 348)

Fig. 14.48

description: Lorica 38-59 * 30-37 urn, cup-shaped, with a smooth inner and serrated outer
collar, slightly indented below apical rim; antapical horn small or lacking; wall reticulate.
Trophont with 15 or 16 adoral membranelles. Macronuclear nodules 2.

distribution: Atlantic sector of the Southern Ocean (Klaas, 1997; Thompson et al, 1999);
North (Daday, 1887) and South Atlantic (Alder, 1999); Barents Sea, Kara Sea (Meunier,
1910; Burkovsky, 1976); Pacific (Yoo & Kim, 1990).

Family Epiplocylididae

Epiplocylis undella (Ostenfeld & Schmidt) Jorgensen (1924: 54, fig. 61)
BASIONYM: Cyttarocylis undella Ostenfeld & Schmidt (1902: 181, fig. 30)
SYNONYM: Ptychocylis undella (Ostenfeld & Schmidt) Brandt (1907: 292)
See Kofoid & Campbell (1929), Sassi & Melo (1986) and Alder (1999) for additional synonyms.

ADDITIONAL REFERENCES: Sassi & Melo (1986: 71), Alder (1999: 51)

Fig. 14.49

description: Lorica 100-157 x 60-75 um, campanulate, slightly dilated below smooth
apical rim; antapical horn short; wall with 2 laminae, reticulate in posterior portion only.

distribution: Atlantic sector of the Southern Ocean (Sassi & Melo, 1986); Indian Ocean,
Pacific, Atlantic, Mediterranean Sea (Jorgensen, 1924); cosmopolitan in tropical and
temperate seas.

Epiplocyloides antarctica Balech (1962b: 39, fig. 88)

SYNONYM: Epiplocylis mira Balech (1958a: 80, pi. 1, figs 11, 12)

ADDITIONAL REFERENCE: Alder (1999: 351)

Fig. 14.50

DESCRIPTION: Lorica 63-74 * 41-55 um, cup-shaped; apical end with an inner and outer
collar; rim sometimes serrated; antapical horn indistinct; wall in posterior portion with small-
meshed reticulate structure. General aspect resembles a very small Cymatocylis.

DISTRIBUTION: Bellingshausen Sea (Balech, 1958a); South Atlantic (Alder, 1999).

375
 CILIATES

Family Ptychocylididae

Ptychocylis ostenfeldi Kofoid & Campbell (1929: 189, fig. 356bis)

See Kofoid & Campbell (1929) for synonymy.

Fig. 14.51

description: Lorica 130-180 * 52-72 urn, goblet-shaped, subapically slightly bulging,

antapically tapering or with a distinct horn; apical rim finely serrated; wall finely reticulate.

distribution: Pacific sector of the Southern Ocean (Mamaeva, 1984, 1986); North Atlantic
(Kofoid & Campbell, 1929).

Ptychocylis wailesi Kofoid & Campbell (1929: 190, fig. 357)

Fig. 14.52

description: Lorica 180-195 * 88-95 urn, cylindrical, subapically and supraequatorially

slightly bulging, antapically obconical and without a horn; apical rim irregularly serrated;
wall reticulate.

distribution: Pacific sector of the Southern Ocean (Mamaeva, 1984, 1986); North Pacific
(Kofoid & Campbell, 1929).

Family Rhabdonellidae

Protorhabdonella curta (Cleve) Jorgensen (1924: 57, fig. 65)

BASIONYM: Cyttarocylis striata f. curta Cleve (1901: 922, fig. 3b)

ADDITIONAL REFERENCE: Alder (1999: 351)

Fig. 14.53

description: Lorica 38-48 * 23-30 urn, obconical, indistinctly indented just below the
slightly outwardly curved apical end; wall with c. 20 delicate and usually longitudinal ribs.

distribution: Weddell Sea (Song & Wilbert, 2000a); South Atlantic, South Indian Ocean
(Cleve, 1901); Pacific (Kofoid & Campbell, 1929; Yoo & Kim, 1990); Mediterranean Sea,
Red Sea (Jorgensen, 1924).

Protorhabdonella simplex (Cleve) Jorgensen (1924: 57, fig. 64)

BASIONYM: Cyttarocylis simplex Cleve (1900a: 972, fig. [7])

ADDITIONAL REFERENCE: Small & Lynn (1985: 444)

Fig. 14.54a-c

description: Lorica 38-70 * 26-38 um, bullet-shaped, indistinctly indented below toothless
apical rim, posteriorly pointed; wall with a few delicate longitudinal ribs. Trophont with
c. 22 somatic kineties (c. 5 shorter and 1 distinctly elongate ciliary rows in lateral field, 7 in

left and 8 in right field, ventral klnety, 1 dorsolateral kinety). Macronuclear nodules 2;
1

micronuclei 2.

distribution: Weddell Sea (Song & Wilbert, 2000a); Atlantic (Cleve, 1900a); Mediterr-
anean Sea, Red Sea, Indian Ocean (Jorgensen, 1924); Pacific (Yoo & Kim, 1990).

Rhabdonella amor (Cleve) Brandt (1907: 327)

BASIONYM: Cyttarocylis amor Cleve (1900a: 970, fig. [4])

SYNONYM: Ptychocylis (Rhabdonella) amor (Cleve) Brandt (1906: 27, pi. 54, figs 4-6)

ADDITIONAL REFERENCES: Laackmann (1910a: 463), Jorgensen (1924: 58)

Fig. 14.55a, b

376
 CILIATES

Figures 14.52-57. Ptychocylis, Protorhabdonella, Rhabdonella, Parapetalotricha and

Cyttarocylis species. 14.52, Ptychocylis wailesi, lorica (from Kofoid & Campbell, 1929). 14.53,
Protorhabdonella curta, lorica (from Cleve, 1901a). 14.54, Protorhabdonella simplex, ventral
(a) and dorsal sides (b) after protargol impregnation (from Small & Lynn, 1985; scale
approximate, originally not provided) and lorica (c, from Cleve, 1900). 14.55, Rhabdonella
amor, lorica shapes (from Laackmann, 1910a). 14.56, Parapetalotricha meridiana, lorica (from
Hada, 1970). 14.57, Cyttarocylis cassis, lorica shapes (from Brandt, 1906). Scale bars: 52 = 40
p.m; 53, 54, 55b = 20 urn; 55a = 30 [im; 56 = 10 urn; 57a = 70 urn; 57b = 50 um. Abbrev-
iations: Dl, dorsolateral kinety; Ic, inner collar; L, lateral field of kineties; Lf, left field of
kineties; Oc, outer collar; Rf, right field of kineties; Vk, ventral kinety.

377
 CILIATES

-Oc

58a

Figure 14.58. Cymatocylis calyciformis. Left side from life (a), left (b) and right sides (c)
after protargol impregnation, and lorica shapes (d, e) (a-d, from Petz et al., 1995; e, from
Kofoid & Campbell, 1929). Scale bars: a, d = 150 um; b, c = 50 urn; e = 60 urn.
Abbreviations: cV, contractile vacuole; Dl, dorsolateral kinety; Ic, inner collar; L, lateral
field of kineties; Lf, left field of kineties; Mn, macronucleus; Oc, outer collar; Rf, right field
of kineties; Vk, ventral kinety.

378
 CILIATES

description: Lorica 80-110 x 40-60 urn, obconical; apical rim toothless; antapical horn
lacking to distinct; wall with delicate and slightly spiralled ribs.

DISTRIBUTION: Weddell Sea (Song & Wilbert, 2000a); tropical and North Atlantic, Indian
Ocean (Brandt, 1907; Laackmann, 1910a); Mediterranean Sea (Jorgensen, 1924).

Family Petalotrichidae

Parapetalotricha meridiana Hada (1970: 38, fig. 59)

Fig. 14.56

description: Lorica c. 30 * 22 urn; wall hyaline, smooth, the short outer collar everted
funnel-like, the very short inner collar straight; bowl obconical.

distribution: Indian sector of the Southern Ocean (Hada, 1970); Bellingshausen Sea
(Kloser, 1990).

Family Cyttarocylididae

Cyttarocylis cassis (Haeckel) Fol (1881: 22, pi. 1, fig. 6)

BASIONYM: Dictyocysta cassis Haeckel (1873: 563, pi. 27, figs 1-3)

SYNONYM: Cyttarocylis conica (Brandt) Kofoid & Campbell (1929: 112, fig. 220)

ADDITIONAL REFERENCES: Laackmann (1910a: 446), Jorgensen (1924: 78), Alder (1999: 351)

Fig. 14.57a, b

description: Lorica 110-215 (-325) x 75-150 urn, broadly to narrowly obconical; short
collar everted funnel-like; antapical horn lacking or indistinct; wall reticulate.

distribution: Atlantic sector of the Southern Ocean (Wasik & Mikolajczyk, 1990); Atlantic,
Indian Ocean (Laackmann, 1910a); Mediterranean Sea (Jorgensen, 1924).

Family Xystonellidae

Cymatocylis calyciformis (Laackmann) Laackmann (1910a: 391, pi. 36, fig. 4; pi. 42,
fig. 12)

BASIONYM: Cyttarocylis calyciformis Laackmann (1907: 236, fig. 3)

SYNONYMS: Cymatocylis brevicaudata (Laackmann) Kofoid & Campbell (1929: 125, fig. 272)
Cymatocylis cylindroides (Laackmann) Kofoid & Campbell (1929: 134, fig. 270)

ADDITIONAL REFERENCES: Van der Spoel (1986: 7), Petz et al. (1995: 151), Alder ( 1999: 349)

Fig. 14.58a-e

DESCRIPTION: Lorica 125-520 x 115-150 um; bowl cup-shaped; inner collar serrated; outer
collar bent downwards; antapical horn highly variable, often long; wall finely reticulate
(appears hyaline at low magnification and in fixed specimens), without agglutinated particles.
Trophont with 41-51 somatic kineties (c. 17 in lateral field, 15 in left and 12 in right field, 1
ventral kinety, 1 dorsolateral kinety). Adoral membranelles 17 external and 1 internal.
Macronuclear nodules 2. Contractile vacuole probably 1. Movement: moderately fast.

distribution: Indian sector of the Southern Ocean (Laackmann, 1907; Hada, 1970);
Bellingshausen Sea (Kloser, 1990); Atlantic sector of the Southern Ocean (Van der Spoel,
1986; Wasik & Mikolajczyk, 1994a; Thompson et al, 1999); Weddell Sea (e.g. Heinbokel &
Coats, 1986; Petz et al, 1995); South Atlantic (Van der Spoel, 1986).

Cymatocylis convallaria Laackmann (1910a: 383, pi. 36, fig. 5; pi. 43, figs 1-4)

SYNONYMS: Cymatocylis affinis Laackmann (1910a: 384, pi. 43, figs 5-15)

379
 CILIATES

Cymatocyhs affinis/convallaria Laackmann 1910a (e.g. Boltovskoy & Alder, 1992; Wasik & Mikolajczyk,
1994b)
lAmphorella antarctica Cleve (1901: 923, fig. 1)

^Cymatocylis antarctica (Cleve) Kofoid & Campbell (1929: 123, fig. 235)
Cyttarocylis parva Laackmann (1907: 237, fig. 5)

Cymatocyhs parva (Laackmann) Laackmann (1910a: 386, pi. 35, fig. 6)

Coxliella cymatiocoides Kofoid & Campbell (1929: 96, fig. 200)

Cymatocyhs gaussi Kofoid & Campbell (1929: 138, fig. 274)
Cymatocyhs scyphus Kofoid & Campbell (1929: 142, fig. 268)
Additional synonyms: see Jorgensen (1924), Petz et a!. (1995) and Alder (1999).

ADDITIONAL REFERENCES: Kofoid & Campbell (1929: 132), Balech (1947: 84), Hada (1961: 144),
Sassi & Melo (1986: 76), Van der Spoel (1986: 7), Barria de Cao (1987: 279), Wasik &
Mikolajczyk (1992: 233, 1994b: 79), Petz et al (1995: 154), Wasik (1998: 10), Alder (1999: 349)

Fig. 14.59a-i

DESCRIPTION: Lorica 55-226 x 46-130 um, inverted bell-shaped; inner collar serrated, wavy
or smooth; outer collar usually bent downwards; antapically blunt to pointed; wall hyaline,
without agglutinated particles. Trophont with 37-39 somatic kineties (c. 15 in lateral field,
12 in left and 9 in right field, 1 ventral kinety, 1 dorsolateral kinety). Adoral membranelles
18 external and 1 internal. Macronuclear nodules 2. Contractile vacuole probably 1.
Movement: moderately fast. Rarely, lorica goblet-shaped or coxlielliform (cup-shaped,
posteriorly narrowly to broadly rounded; wall annulated).

distribution: Indian sector of the Southern Ocean (Laackmann, 1910a; Hada, 1970); Pacific
sector of the Southern Ocean (Balech, 1958b; Tumantseva, 1989); Ross Sea (Hopkins, 1987;
Monti & Fonda Umani, 2000); Bellingshausen Sea (e.g. Balech, 1958a, 1973a; Barria de
Cao, 1987; Kloser, 1990); Atlantic sector of the Southern Ocean (e.g. Wasik & Mikolajczyk,
1990; Thompson et al, 1999); Weddell Sea (e.g. Balech & El-Sayed, 1965; Boltovskoy &
Alder, 1992; Petz et al, 1995); South Atlantic (Alder, 1999); South Indian Ocean (Cleve,
1901); North Pacific (Kofoid & Campbell, 1929).

Boltovskoy & Alder (1992) suggested conspecificity of C. convallaria with the much larger
C. drygalskii, whereas Williams et al. (1994) maintained C. convallaria and C. parva as
separate species.

Cymatocylis cristallina Laackmann (1910a: 374, pi. 40, figs 1-5)

SYNONYMS: Cymatocyhs crassa Kofoid & Campbell (1929: 132, fig. 254)
Cymatocyhs culcullus Kofoid & Campbell (1929: 133, fig. 261)
Cymatocyhs glans Kofoid & Campbell (1929: 139, fig. 269)
Cymatocyhs subconica Kofoid & Campbell (1929: 144, fig. 238)
For additional synonyms, see Alder (1999).

ADDITIONAL REFERENCE: Alder (1999: 349)

Fig. 14.60a-c

description: Lorica 180-260 * 100-120 urn, chalice-shaped to obconical or cylindrical; rim

everted; inner collar serrated; sometimes with a very short antapical horn; wall hyaline, with
indistinct irregularly distributed striae.

distribution: Indian sector of the Southern Ocean (Laackmann, 1910a; Hada, 1970); Ross
Sea (Monti & Fonda Umani, 1995, 2000); Atlantic sector of the Southern Ocean (Thompson
et al, 1999); Weddell Sea (Heinbokel & Coats, 1986); South Atlantic (Alder, 1999).

Cymatocylis drygalskii (Laackmann) Laackmann (1910a: 376, pi. 41, figs 1-8)

basionym: Cyttarocylis drygalskii Laackmann (1907: 236, fig. 2)

SYNONYMS: Cymatocylis flava f. typica Laackmann (1910a: 371, pi. 39, figs 2, 4), pro parte
Cymatocylis cylindrus (Laackmann) Kofoid & Campbell (1929: 135, fig. 259)

380
 CILIATES

Figure 14.59a-e. Cymatocylis convallaria. Right side from life (a), right ventrolateral (b)
and left dorsolateral (c) sides after protargol impregnation, and typical (d) and ptycho-
cyliform (e) loricae (from Petz et al, 1995). Scale bars: a-e = 50 um. Abbreviations: Dl,
dorsolateral kinety; eAM, elongate external adoral membranelles; Fv, food vacuole; iAM,
internal adoral membranelle; L, lateral field of kineties; Lf, left field of kineties; O, oral
cavity; Pf, pharyngeal fibres; pM, paroral membrane; Rf, right field of kineties; Vk, ventral
kinety.

381
 CILIATES

Figures 14.59f-64. Cymatocylis species. 14.59f-i, C. convallaria, lorica shapes (f, as

C. affinis, C. gaussi; g, as C. antarctica; h, as C. parva; i, as Coxliella cymatiocoides) (f, from
Laackmann, 1910a; from Kofoid & Campbell, 1929; h, from Laackmann, 1907). 14.60,
g, i,

(from Laackmann, 1910a). 14.61, C. drygalskii, lorica shapes (a,

C. cristallina, lorica shapes
from Laackmann, 1907; b, from Laackmann 1910a). 14.62, C. kerguelensis, lorica (from
Laackmann, 1910a). 14.63, C. nobilis, lorica shapes (a, from Laackmann, 1907; b, from
Laackmann, 1910a). 14.64, C. vanhoeffeni, lorica shapes (a, from Laackmann, 1907; b, as
C.flava, c, from Laackmann, 1910a). Scale bars: 59f, = 40 urn; 59g, h, 62 = 20 urn; 60a, b =
i

80 pm; 60c, 64b = 60 pm; 61, 63, 64c = 50 pm; 64a = 100 pm.

382
 CILIATES

Cymatocylis ecaudata Kofoid & Campbell (1929: 137, fig. 263)

Cymatocylis ovata (Laackmann) Kofoid & Campbell (1929: 141, fig. 257)
Cymatocylis typica Laackmann, in Kofoid & Campbell (1929: 145, fig. 260)
Additional synonyms: see Alder (1999), who also considered Co.xliella intermedia to be conspecific with C.
drygalskii; see also notes under C. convallaria.

ADDITIONAL REFERENCES: Balech (1947: 81), Sassi & Melo (1986: 76), Van der Spoel (1986: 7),
Barria de Cao (1987: 278), Alder (1999: 349)

Fig. 14.61a, b

description: Lorica 140-340 x 70-125 um, cylindrical; rim serrated and bent downwards,
often with a short antapical horn; wall hyaline. Coxlielliform lorica cylindrical, annulated.
Trophont with 19 adoral membranelles. Macronuclear nodules 2; micronuclei 2.

distribution: Indian sector of the Southern Ocean (Laackmann, 1910a; Grey et al, 1997);
Ocean (Balech, 1958b); Ross Sea (e.g. Monti & Fonda Umani,
Pacific sector of the Southern
1995; Fonda Umani et al, 1998); Bellingshausen Sea (e.g. Balech 1958a, 1973a; Barria de
Cao, 1987); Atlantic sector of the Southern Ocean (e.g. Wasik & Mikolajczyk, 1990;
Thompson et al, 1999); Weddell Sea (e.g. Balech & El-Sayed, 1965; Heinbokel & Coats,
1986; Boltovskoy & Alder, 1992); South Atlantic (Alder, 1999).

Cymatocylis kerguelensis Laackmann (1910a: 387, pi. 35, fig. 5)

See Alder (1999) for synonymy.

additional reference: Alder (1999: 349)

Fig. 14.62

description: Lorica 55-67 * 50-65 urn, sac-shaped; inner collar serrated; outer collar bent
outwards and up; posteriorly broadly rounded, without an antapical horn; wall distinctly
striated.

distribution: Indian sector of the Southern Ocean (Laackmann, 1910a); Bellingshausen Sea
(Kloser, 1990); Atlantic sector of the Southern Ocean (Wasik & Mikolajczyk, 1994a); South
Atlantic (Alder, 1999); South Indian Ocean (Hada, 1970).

Cymatocylis nobilis (Laackmann) Laackmann (1910a: 389, pi. 42, figs 1-6)

BASIONYM: Cyttarocylis nobilis Laackmann (1907: 237, fig. 4)

SYNONYM: Cymatocylis folliculus Kofoid & Campbell (1929: 138, fig. 264)
For additional synonyms, see Alder (1999).

ADDITIONAL REFERENCES: Balech (1958b: 399), Van der Spoel (1986: 7), Alder (1999: 349)

Fig. 14.63a, b

DESCRIPTION: Lorica 175^150 x 100-160 urn; bowl cylindrical and long; inner collar
serrated; outer collar bent downwards; antapical horn usually long; wall hyaline.
distribution: Indian sector of the Southern Ocean (Laackmann, 1907); Pacific sector of the
Southern Ocean (Balech, 1958b); Ross Sea (Monti & Fonda Umani, 1995, 2000);
Bellingshausen Sea (Kloser, 1990); Atlantic sector of the Southern Ocean (e.g. Van der
Spoel, 1986; Wasik & Mikolajczyk, 1990; Thompson et al, 1999); South Atlantic (Alder,
1999).

Cymatocylis vanhoeffeni (Laackmann) Laackmann (1910a: 358, pi. 33; pi. 36, fig. 1)

BASIONYM: Ptychocylis vanhoeffeni Laackmann (1907: 239, fig. 9)

SYNONYMS: Cymatocylis flava Laackmann (1910a: 369, pi. 39, figs 1, 3, 5-10), pro parte
Cymatocylis calyx Kofoid & Campbell (1929: 129, fig. 246)
Cymatocylis conica (Laackmann) Kofoid & Campbell (1929: 131, fig. 245)
Cymatocylis digitulus Kofoid & Campbell (1929: 136, fig. 234)

383
 CILIATES

Protocymatocylis subrotundata (Laackmann) Kofoid & Campbell (1929: 120, fig. 229)
Cymatocylis subrotundata (Laackmann) Balech (1973a: 31, figs 165-174)
Additional synonyms: see Alder (1999), who also considered Coxliella frigida to be a synonym of
C. vanhoeffeni.

additional REFERENCES: Laackmann (1910a: 358), Balech (1947: 83, 1958b: 400), Sassi &
Melo (1986: 77), Van der Spoel (1986: 7), Barria de Cao (1987: 279), Alder (1999: 349)
Figs 14.64a-c; 14.191

DESCRIPTION: Lorica 240-600 x 82-130 [tm; bowl elongate, chalice-shaped to obconical,

apically everted; rim serrated, without an outer collar; antapical horn usually long,
sometimes lacking; wall hyaline, striated (particularly in anterior portion of lorica),
appearing brownish. Coxlielliform lorica cylindrical, annulated, apically everted, posteriorly
rounded, without an antapical horn.

DISTRIBUTION: Indian sector of the Southern Ocean (Laackmann, 1907; Hada, 1970); Pacific
sector of the Southern Ocean (Balech, 1958b; Tumantseva, 1989); Ross Sea (e.g. Littlepage,
1968; Monti & Fonda Umani, 1995); Bellingshausen Sea (e.g. Balech, 1973a; Kloser, 1990;
Alder & Boltovskoy, 1991a); Atlantic sector of the Southern Ocean (e.g. Sassi & Melo,
1986; Wasik & Mikolajczyk, 1990; Thompson et al, 1999); Weddell Sea (e.g. Balech &
El-Sayed, 1965; Heinbokel & Coats, 1986; Boltovskoy & Alder, 1992; Petz et al, 1995);
South Atlantic (Alder, 1999).

Protocymatocylis pseudoconica Hada (1961: 143, fig. 3)

ADDITIONAL REFERENCE: Hada (1970: 34)

Fig. 14.65

DESCRIPTION: Lorica 55-150 x 15-40 um, elongate-obconical; wall with numerous short
longitudinal ridges.

distribution: Indian sector of the Southern Ocean (Hada, 1970); Pacific sector of the
Southern Ocean (Hada, 1961).

Xystonella acus (Brandt) Laackmann (1910a: 451)

basionym: Cyttarocylis (Xystonella) acus Brandt (1906: 6, pi. 45, figs 4-6)

ADDITIONAL REFERENCES: Brandt (1907: 244), Jorgensen (1924: 34)

Fig. 14.66a, b

description: Lorica 250-580 * 50-115 um, elongate chalice-shaped, apically usually

slightly dilated, frequently somewhat widened in middle portion; antapical horn highly
refractile, thus easily recognisable, of variable length; wall hyaline and reticulate. Trophont
with 2 macronuclear nodules and 2 micronuclei.

distribution: Weddell Sea (Song & Wilbert, 2000a); Atlantic (Brandt, 1907; Laackmann,
1910a); Mediterranean Sea (Jorgensen, 1924).

Family Undellidae

Undella claparedii (Entz) Daday (1887: 566, pi. 19, fig. 1)

BASIONYM: Tintinnus claparedii Entz (1885: 202, pi. 14, figs 10, 11)
SYNONYMS: Proplectella claparedii (Entz) Kofoid & Campbell (1929: 276, fig. 525)

Continued ... 14.75, Salpingella acuminatoides, lorica (from Laackmann, 1910a). 14.76,
Salpingella costata, lorica (from Laackmann, 1910a). 14.77, Salpingella decurtata, lorica
(from Jorgensen, 1924). Scale bars: 65, 74, 76, 77 = 30 um; 66 = 100 um; 67, 71, 72 = 20
um; 68 = 60 um; 69 = 40 um; 70 = 10 um; 73, 75 = 50 um. Abbreviations: K, knob; R, ribs.

384
 CILIATES

Figures 14.65-77. Protocymatocylis, Xystonella, Amphorellopsis, Amphorides,

Undella,
Bursaopsis, Dadayiella, Daturella and Salpingella species.
14.65, Protocymatocylis
pseudoconica, lorica (from Hada, 1970). 14.66, Xystonella acus, lorica shapes (from Brandt,
1906). 14.67, Undella claparedii, lorica (from Brandt, 1907). 14.68, Undella hyalina, lorica
(from Kofoid & Campbell, 1929). 14.69, Amphorellopsis quinquealata, lorica (from
Laackmann, 1907). 14.70, Amphorides laackmanni, lorica (as Tintinnus amphora var. dadayi,
from Laackmann, 1910a). 14.71, Bursaopsis conicoides, lorica (from Hada, 1970). 14.72,
Bursaopsis ollula, lorica (from Hada, 1970). 14.73, Dadayiella ganymedes, lorica shapes (b, as
Tintinnus bulbosus) (from Brandt, 1906). 14.74, Daturella frigida, lorica (from Hada, 1970).

385
 CILIATES

Figures 14.78, 79. Pelagostrobilidium and Rimostrombidium species. 14.78, P. neptuni, right
side from life (a) and antapical side (b) after protargol impregnation (from Petz et al., 1995).
14.79, R. conicum, left ventrolateral side from life (a) and dorsal (b) and right (c) sides of
very early divider after protargol impregnation (from Agatha & Riedel-Lorje, 1998). Scale
bars: 78a = 30 pm; 78b = 20 pm; 79 = 10 pm. Abbreviations: cV, contractile vacuole; eAM,
external adoral membranelles; Kl-5, somatic kineties 1-5; Mi, micronucleus; Mn, macro-
nucleus; Op, oral primordium; Sk, somatic kinety.

386
 CILIATES

Proplectella globosa (Brandt) Kofoid & Campbell (1929: 278, fig. 541)
Undella antarctica Hada (1970: 39, fig. 60)
For additional synonyms, see Balech (1959) and Alder (1999).

ADDITIONAL REFERENCES: Brandt (1907: 362), Laackmann (1910a: 471), Jorgensen (1924: 38),
Balech (1959: 47), Alder (1999: 350)

Fig. 14.67

description: Lorica 53-95 x 38-72, sac-like; inner collar slightly funnel-shaped; subapical
margin indistinctly inflated (with the wall thickened), antapically broadly rounded; wall
hyaline. Trophont with 2 macronuclear nodules and 4 micronuclei.

distribution: Indian sector of the Southern Ocean (Hada, 1970); Indian Ocean, Pacific
& Cunha, 1917); Atlantic, Mediterranean Sea (Daday,
(Brandt, 1907); South Atlantic (Faria
1887).

Undella hyalina Daday (1887: 564, pi. 18, fig. 17)

SYNONYMS: Undella declivis Kofoid & Campbell (1929: 261, fig. 507)
Undella parva Kofoid & Campbell (1929: 264, fig. 508)

ADDITIONAL REFERENCES: Jorgensen (1924: 42), Balech (1959: 46), Alder (1999: 350)

Fig. 14.68

description: Lorica 145-280 * 54-74 um, cylindrical, posteriorly tapering to rounded; wall
hyaline and smooth. Trophont with 2 macronuclear nodules, 2 micronuclei and 2 contractile
vacuoles.

distribution: Atlantic sector of the Southern Ocean (e.g. Wasik & Mikolajczyk, 1990);
South Atlantic (Alder, 1999); tropical Pacific, Atlantic, Mediterranean Sea (Kofoid &
Campbell, 1929); Indian Ocean (Jorgensen, 1924).

Family Tintinnidae

Amphorellopsis quinquealata (Laackmann) Balech (1971: 180, figs 758, 759)

BASIONYM: Tintinnus quinquealatus Laackmann (1907: 236, fig. 1)

SYNONYM: Bursaopsis quinquealata (Laackmann) Kofoid & Campbell (1929: 305, fig. 578)

ADDITIONAL REFERENCES: Laackmann (1910a: 412), Alder (1999: 347)

Fig. 14.69

DESCRIPTION: Lorica 180-215 x 45-57 pm, elongate chalice-shaped; fins 5, longitudinal or

slightly spiralling along entire lorica. Trophont with 2 macronuclear nodules and 2 micro-
nuclei.

distribution: Indian sector of the Southern Ocean (Laackmann, 1907); Bellingshausen Sea
(Balech, 1973a); Weddell Sea (Heinbokel & Coats, 1986); South Atlantic (Alder, 1999).

Amphorides laackmanni (Jorgensen) Strand (1928: 30)

BASIONYM: Amphorella laackmanni Jorgensen (1924: 19, fig. 14)

See Kofoid & Campbell (1929) for synonymy.

ADDITIONAL REFERENCE: Sassi & Melo (1986: 77)

Fig. 14.70

description: Lorica 54-73 x 15-20 pm, elongate, apically indistinctly everted, subequat-
orially slightly inflated, antapically tapering, with few longitudinal fins along entire lorica;
wall hyaline.

387
 CILIATES

distribution: Atlantic sector of the Southern Ocean (Sassi & Melo, 1986); South Atlantic
(Laackmann, 1910a); Mediterranean Sea (Jorgensen, 1924).

Bursaopsis conicoides Hada (1970: 40, fig. 63)

Fig. 14.71

DESCRIPTION: Lorica 25-65 * 17-30 um, obconical; lateral margins straight.

distribution: Indian sector of the Southern Ocean, South Atlantic (Hada, 1970).

A poorly described species.

Bursaopsis ollula Hada (1970: 39, fig. 62)

Fig. 14.72

description: Lorica 15-24 * 12-25 um, obovoid to heart-shaped; apical opening small,
without a collar; wall hyaline.

distribution: Indian sector of the Southern Ocean (Hada, 1970).

A poorly described species.

Dadayiella ganymedes (Entz) Kofoid & Campbell (1929: 321, fig. 610)
BASIONYM: Tintinnus ganymedes Entz (1884: 409, pi. 24, figs 17, 18)

See Alder (1999) for synonymy.

ADDITIONAL REFERENCES: Jorgensen (1924: 22), Balech (1959: 50), Sassi & Melo (1986: 78),
Alder (1999: 346)

Fig. 14.73a, b

description: Lorica 95-133 * 27-38 um, elongate chalice-shaped, apically slightly dilated;
collar with short longitudinal ribs (facets) that sometimes project anteriorly as hair-like
appendages; posteriorly narrowed; antapical horn with few ribs, occasionally with a small
terminal knob; wall hyaline. Trophont with 1 macronucleus and 1 micronucleus.

distribution: Atlantic sector of the Southern Ocean (Sassi & Melo, 1986; Thompson et ah,
1999); cosmopolitan in tropical seas.

Daturella frigida Hada (1970: 40, fig. 65)

Fig. 14.74

description: Lorica c. 100 x 28 um, tubular, anteriorly slightly widened, with a smooth
apical rim, posteriorly narrowing and curved, antapically open, with irregular longitudinal
fins along almost entire lorica; wall hyaline.

distribution: Indian sector of the Southern Ocean (Hada, 1970).

A poorly described species.

Salpingella acuminatoides (Laackmann) Kofoid & Campbell (1929: 350, fig. 673)
BASIONYM: Tintinnus acuminatoides Laackmann (1910a: 409, 480, pi. 50, figs 6-8)
SYNONYMS: Tintinnus acuminatus var. secata ? sensu Laackmann (1907: 235)
Salpingella laackmanni Kofoid & Campbell (1929: 353, fig. 670)
Salpingella sp. aff.faurei sensu Barria de Cao (1987: 280, fig. 3e)

ADDITIONAL REFERENCES: Balech & El-Sayed (1965: 123), Alder (1999: 347)

Fig. 14.75

description: Lorica (1 20—) 230-250 * 10-15 um, nail-shaped, apically widened and funnel-
like, with 4 slightly spiralling fins along posterior portion only, antapically open.

388
 CILIATES

distribution: Bellingshausen Sea (Kloser, 1990); Weddell Sea (Balech & El-Sayed, 1965;
Barria de Cao, 1987); South Atlantic (Alder, 1999).

Salpingella costata (Laackmann) Kofoid & Campbell (1929: 351, fig. 680)
BASIONYM: Tintinnus costatus Laackmann (1910a: 412, pi. 50, figs 9, 10)

ADDITIONAL REFERENCE: Alder (1999: 347)

Fig. 14.76

DESCRIPTION: Lorica 150-215 x 12-20 urn, tubular, apically slightly widened, middle portion
somewhat bulging, posteriorly tapering, antapically open, with distinct longitudinal fins
along entire lorica, these occasionally terminating before antapical pole.

distribution: Indian sector of the Southern Ocean (Laackmann, 1910a); South Atlantic
(Alder, 1999).

Salpingella decurtata Jorgensen (1924: 15, fig. 10)

Fig. 14.77

description: Lorica 120-150 * 16-23 um, nail-shaped, apically indistinctly everted, with
parallel sides, posteriorly tapering, antapically open; with a few slightly spiralling fins along
posterior 25-33% of lorica.
distribution: Weddell Sea (Song & Wilbert, 2000a); Mediterranean Sea (Jorgensen, 1924).

Order Choreotrichida

Family Strobilidiidae

Pelagostrobilidium neptuni (Montagnes & Taylor) Petz, Song & Wilbert (1995: 139,
fig. 41)
BASIONYM: Strobilidium neptuni Montagnes & Taylor (1994: 572, figs 1-3)

Fig. 14.78a, b

description: In vivo 35-60 um diameter. Body bowl-shaped, posteriorly broadly rounded.

Somatic kineties 5; kineties 1 and 3-5 ±longitudinal; kinety 2 forms an arc on posterior left-
lateral side beginning and ending at kinety 1. Adoral zone of membranelles composed of 36-40
external and 1 or 2 internal membranelles in a closed spiral. Macronucleus horseshoe-
shaped; micronuclei 2. Movement: very fast, sometimes jumping, remains stationary in-
between and rotating about main body axis, appears helicopter-like in top view.

distribution: Ross Sea, in the region 68.98°-76.60°S and 162.43°-164.85°W, in the plankton
(Petz, unpublished data); Weddell Sea (Petz et al, 1995); North Pacific (Montagnes &
Taylor, 1994).

Pelagostrobilidium spirale (Leegaard) Petz, Song & Wilbert (1995: 141)

BASIONYM: Lohmanniella spiralis Leegaard (1915: 27, fig. 18)

SYNONYM: Strobilidium spirale (Leegaard) Lynn & Montagnes (1988: 646, figs 3, 4F, G, 5)

In the absence of morphological details, the population tentatively identified by Leakey et al.
(1994) from the Atlantic sector of the Southern Ocean is considered to belong to P. neptuni.
Pelagostrobilidium spirale differs from P. neptuni in having only 1 micronucleus.

389
 CILIATES

Rimostrombidium conicum (Kahl) Petz & Foissner (1992: 160)

BASIONYM: Strobilidium conicum Kahl (1932: 512, fig. 82.33)

ADDITIONAL REFERENCE: Agatha & Riedel-Lorje (1998: 13)

Fig. 14.79a-c

description: In vivo 25-50 x 15-30 um. Body obconical, with indistinct longitudinal
furrows, posteriorly pointed.Cytoplasm reddish to brownish. Somatic kineties 6,
longitudinal and slightly oblique. Adoral membranelles 20-26 external and 1 internal, in a
closed spiral. Macronucleus horseshoe-shaped; micronucleus 1. Contractile vacuole terminal.
Movement: very fast over long distances, sometimes swimming slowly and rotating about
main body axis.

distribution: Indian sector of the Southern Ocean (Hada, 1970), but obviously
misidentified; North Sea (Kahl, 1932).

Rimostrombidium glacicolum Petz, Song & Wilbert (1995: 141, fig. 42)

Figs 14.80a-c; 14.178

description: In vivo 30-45 x 20-30 urn. Body turbinate. Somatic kineties 5, short,
longitudinally arranged in posterior third of body, covered by a cytoplasmic flap. Adoral
membranelles 17-20, all external in a closed spiral. Macronucleus reniform; micronucleus 1.
Movement: moderately fast.
distribution: Ross Sea, in the region 68.80°-76.02°S and 164.12°E-134.88°W, in sea-ice
and plankton under ice (Petz, unpublished data; Petz & Valbonesi, unpublished data);
Weddell Sea (Petz et al, 1995).

Family Leegaardiellidae

Leegaardiella elbraechteri Petz (1995: 134, fig. 40)

Fig. 14.81a-c

DESCRIPTION: In vivo 50-65 x 40-55 urn. Body cornucopia-shaped, covered by a posteriorly

open hyaline 'lorica'. Somatic kineties 4, short, longitudinally radiating from near posterior
pole, without a cytoplasmic flap. Adoral membranelles 19-21 external, continuous with 4 or
5 distinct internal membranelles in a closed spiral; rightmost internal membranelle enlarged.
Macronuclear nodules 2; micronuclei 2. Contractile vacuole in posterior third of body.
Movement: slow to moderately fast, rarely very rapid, rotating about main body axis, often
swimming backwards. Secretes a mucous thread at antapical end when undisturbed and
attaches to substratum, remains rotating on the spot; disturbed individual immediately
detaches from thread and darts away.

distribution: Ross Sea, in the region 68.98°-76.60°S and 1 62.43°- 1 64.85° W, in surface
plankton (Petz, unpublished data); Weddell Sea (Petz et al, 1995).

Family Lohmanniellidae

Lohmanniella oviformis Leegaard (1915: 28, fig. 19)

ADDITIONAL REFERENCES: Kahl (1932: 513), Lynn & Montagnes (1988: 649)

Fig. 14.82a. b

description: Fixed specimens 12-44 x 11-49 urn. Body subspherical. Somatic kineties 5,
very short, radiating star-like from posterior pole, composed of non-ciliated monokinetids.
Adoral membranelles 17-21 external and 4-6 internal in a closed spiral. Macronucleus
reniform to globular.

390
 CILIATES

AZM

80a

81a

Figures 14.80, 81. Rimostrombidium and Leegaardiella species. 14.80, R. glacicolum, dorsal
side from life (a) and ventral (b) and dorsal (c) sides after protargol impregnation (from Petz
et al, 1995). 14.81, L. elbraechteri, left side from life (a, arrowhead points to enlarged
internal adoral membranelle), and ventral (b) and dorsal (c) sides after protargol impreg-
nation (from Petz et al, 1995). Scale bars: 80 = 10 urn; 81 = 30 um. Abbreviations: AZM,
adoral zone of membranelles; cV, contractile vacuole; Fv, food vacuole; iAM, enlarged
internal adoral membranelle; L, lorica; M, mucous thread; Sk, somatic kinety.

391
 CILIATES

83a

Figures 14.82, 83. Lohmanniella and Notocephalus species. 14.82, L. oviformis, lateral (a)
and posterior (b) sides after protargol impregnation (from Lynn & Montagnes, 1988). 14.83,
N. parvulus, ventral side from life (a) and ventral (b) and dorsal (c) sides after protargol
impregnation (from Petz et ai, 1995). Scale bars: 82 = 10 urn; 83 = 50 urn Abbreviations:
AZM, adoral zone of membranelles; DK, dorsal kineties; FC, frontal cirri; MC, mid-ventral
cirri; RMC, right-marginal cirri; SK, somatic kinety; TC, transverse cirri.

392
 CILIATES

nc^y

Figures 14.84-86. Metaurostyla, Metaurostylopsis and Holosticha species. 14.84, Metaur-

ostyla thompsoni, ventral side after silver nitrate impregnation (from Thompson, 1972).
14.85, Metaurostylopsis rubra, ventral side from and ventral (b) and dorsal (c) sides
life (a),
after protargol impregnation (from Song & Wilbert, 2002). 14.86, Holosticha foissneri,
ventral side from life (a), and ventral (b) and dorsal (c) sides after protargol impregnation
(from Petz et al, 1995). Scale bars: 84 = 20 pm; 85 = 30 pm; 86 = 40 pm. Abbreviations:
AZM, adoral zone of membranelles; BC, buccal cirrus; FC, frontal cirri; FTC, fronto-
terminal cirri; LMC, left-marginal cirral rows; Mi, micronucleus; Mn, macronucleus; MVC,
midventral cirri.

393
 CILIATES

87a

Figures 14.87, 88. Holosticha species. 14.87, H. pullaster, ventral side from life (a), and
ventral (b) and dorsal (c) sides after protargol impregnation (from Petz et al., 1995). 14.88,
H. spindleri, ventral side from life (a), and ventral (b) and dorsal (c) sides after protargol
impregnation (from Petz et al., 1995). Scale bars: 87 = 20 urn; 88 = 40 urn. Abbreviations:
cV, contractile vacuole; FTC, frontoterminal cirri; pAM, posteriormost elongated adoral
membranelle.

394
 CILIATES

distribution: Indian sector of the Southern Ocean (Grey et al., 1997); North Atlantic, North
Sea (Leegaard, 1915; Lynn & Montagnes, 1988); probably cosmopolitan.

Order Urostylida

Family Urostylidae

Holosticha foissneri Petz, Song & Wilbert (1995: 159, fig. 47)

Fig. 14.86a-c

DESCRIPTION: In vivo 120-190 * 35-60 um. Body elongate-ellipsoidal to slightly fusiform,

dorsoventrally flattened. Cirri: 3 frontal, 1 buccal, 2 or 3 frontoterminal, 9-17 transverse.
Midventral row of 16-20 oblique cirral pairs extending to transverse cirri. Marginal cirral
rows 1 right, 1 left, the latter obliquely bent at anterior end. Dorsal kineties 4. Adoral
membranelles 26-36 (anteriorly, 7-13 membranelles slightly separate). Macronuclear nodules
(5-) 8 (-1 1); micronuclei 2-6. Contractile vacuole subequatorial. Movement: crawling slowly.

distribution: Ross Sea, in the region 69.97°-71.03°S and 135.07°-165.02°W, in sea-ice and
plankton (Petz, unpublished data); Weddell Sea (Petz et al., 1995).

Holosticha pullaster (Muller) Foissner, Blatterer, Berger & Kohmann (1991: 240, figs
1-14)

BASIONYM: Richoda pullaster Muller (1773: 81)

SYNONYM: Holosticha diademata (Rees) Kahl sensu Song & Wilbert (2002: 53)
For additional synonyms, see Foissner el al. (1991).

ADDITIONAL REFERENCE: Petz et al. (1995: 166)

Fig. 14.87a-c

DESCRIPTION: In vivo 50-100 * um. Body elongate-ellipsoidal, dorsoventrally

20-35
flattened. Cirri: 3 frontal, 1 buccal, 2 frontoterminal, 6-8 transverse. Midventral
row of 10 or
11 cirral pairs extending to transverse cirri. Marginal cirral rows 1 right, 1 left, the latter
obliquely bent at anterior end. Dorsal kineties 4 or 5. Adoral membranelles 10-28
(anteriorly, 3-8 membranelles slightly separate). Macronuclear nodules 2; micronuclei 2.
Contractile vacuole distinctly subequatorial. Movement: crawling slowly, sometimes jerking
back and forth, resting for long periods.

distribution: Ross Sea, in the region 65.28°-74.03°S and 134.85°-165.42°W, in sea-ice

(Petz, unpublished data); Atlantic sector of the Southern Ocean (Song & Wilbert, 2002);
Weddell Sea (Petz et al., 1995); widespread in freshwater (Foissner et al., 1991), probably
cosmopolitan.

Often confused with H. diademata (Rees) Kahl, which has the contractile vacuole in mid-
body (Foissner et al., 1991).

Holosticha spindleri Petz, Song & Wilbert (1995: 164, figs 48, 59, 60)

Figs 14.88a-c; 14.175; 14.176

description: In vivo 100-200 * 45-75 um. Body ellipsoidal to broadly fusiform,

dorsoventrally flattened. Cirri: 3 frontal, 1 buccal, 2 frontoterminal, 7-14 transverse.
Midventral row of 9-17 oblique cirral pairs extending to transverse cirri. Marginal cirral rows
1 right, 1 left, the latter obliquely bent at anterior end. Dorsal kineties 4 or 5. Adoral
membranelles 22-A2> (anteriorly, 4-18 membranelles slightly separate), posteriormost
membranelle c. twice as long as adjacent ones. Macronuclear nodules (1-) 4; micronuclei 2-4.
Contractile vacuole equatorial. Movement: crawling.

395
 CILIATES

distribution: Ross Sea, in the region 69.97°-73.60°S and 134.85°-165.03°W, in sea-ice and
rarely in the plankton (Petz, unpublished data); Weddell Sea (Petz et al., 1995).

Metaurostyla thompsoni Jankowski (1979: 84)

SYNONYM: Urostyla sp. (Thompson, 1972: 285, fig. 26)

Fig. 14.84

description: Fixed specimens 103-150 * 52-75 urn. Body ellipsoidal, dorsoventrally

flattened. Cirri: 3 (or 4) frontal, 1 buccal, 6 frontoterminal, 3-5 transverse. Midventral row
posteriorly shortened, composed of c. 9 cirral pairs in anterior and c. 7 single cirri in
posterior portion. Marginal cirral rows c. 4 or 5 right and 5 or 6 left. Dorsal kineties 7.
Adoral membranelles c. 40. Macronuclear nodules several.

DISTRIBUTION: Bellingshausen Sea (Thompson, 1972).

Metaurostylopsis rubra Song & Wilbert (2002: 52, figs 12, 49, 50)

Fig. 14.85a-c

DESCRIPTION: In vivo 150-300 * 50-90 um. Body elongate-ellipsoidal, flexible, contractile,

with brick-red pigments in cytoplasm; dorsoventrally flattened. Cortical granules on dorsal
side (colour unknown). Cirri: 4 frontal, buccal, 5-8 frontoterminal, 4-6 transverse.
1

Midventral row composed of 8-11 cirral pairs in anterior, 8-13 single cirri in posterior
portion. Marginal cirral rows 6 or 7 right, 6-9 left. Dorsal kineties 3 long and 2-4 short.
Adoral membranelles 35-46. Macronuclear nodules c. 100. Contractile vacuole supra-
equatorial. Movement: crawling moderately quickly.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002).

Notocephalus parvulus (Corliss & Snyder) Petz, Song & Wilbert (1995: 170, fig. 50)
BASIONYM: Tachysoma parvulum Corliss & Snyder (1986: 44, fig. 6)

Fig. 14.83a-c

description: In vivo 155-180 x 40-45 um. Body fusiform, frontal portion enlarged, head-
like, posteriorly tapering, dorsoventrally oral cavity deep, with a
slightly flattened;
conspicuous cytoplasmic lip on right. Cirri: 3 frontal and 5-7 transverse. Marginal cirral
rows 1 right and left. Midventral row shortened, composed of 5-9 cirral pairs (sometimes
1

irregular). Dorsal kineties 3. Adoral zone of c. 55-58 membranelles spoon-shaped, extends

far onto right side. Macronuclear nodules 2; micronuclei (1 or) 2. Contractile vacuole supra-
equatorial. Movement: slowly crawling.

distribution: Ross Sea, at 73°36'S 134°51'W, in nilas ice (Petz, unpublished data); Weddell
Sea (Corliss & Snyder, 1986; Petz et al., 1995).

Family Pseudokeronopsidae

Thigmokeronopsis antarctica Petz (1995: 138, figs 1-4)

Fig. 14.89a-c

description: In vivo c. 160-380 * 50-115 um. Body elongate, posteriorly tapering,

dorsoventrally slightly flattened; buccal Field large; cytoplasm brownish red. Cirri: 20-42
frontal, in 2 rows, forming a bicorona, continuous with short midventral row of c. 40-63 cirral
pairs and, sometimes, a few single cirri posteriorly; 1-3 frontoterminal, 2-5 buccal, no
row of Fine cirri
distinct transverse cirri. Left postoral ventral (sometimes with a very short 1

second row); marginal cirral rows right and, usually,

1 left. Dorsal kineties 3 (or 4). Adoral
1

zone of 45-71 membranelles extending far onto right side. Macronuclear nodules c. 120-170;
micronuclei 2-5. Contractile vacuole supraequatorial. Movement: crawling slowly.

distribution: Weddell Sea (Petz, 1995).

396
 CILIATES

wmm

90a

Figures 14.89-91. Thigmokeronopsis and Laboea species. 14.89, T. antarctica, ventral side
from life (a), and ventral (b) and dorsal (c) sides after protargol impregnation (from Petz,
1995). 14.90, T. crystallis, ventral side from life (a), and ventral (b) and dorsal (c) sides after
protargol impregnation (from Petz, 1995). 14.91, L. strobila, ventral sides after fixation (a,
after Wulff, 1919) and protargol impregnation (b, from Montagnes et ai, 1988). Scale bars:
89a, 90b, c = 50 urn; 89b, c = 100 am; 90a = 40 am; 91 = 30 urn. Abbreviations: aAM,
anterior adoral membranelles; B, bicorona; Bf, baccal field; C, cytopyge; DK, dorsal kinety;
Ek, equatorial kinety; eM, endoral membrane; Lp, left postoral ventral cirral row; Mi,
micronucleus; Mn, macronuclear nodule; pM, paroral membrane; RMC, right-marginal cirri;
S, subpellicular platelet layer; TC, transverse cirri; vAM, ventral adoral membranelles.

397
 CILIATES

Figure 14.92. Tontonia antarctica. Ventral side of trunk (a) and specimen with tail extended
(b) from life, and ventral (c) and dorsal (d) sides after protargol impregnation (tail
contracted) (from Petz et ai, 1995). Scale bars: 92a. c, d = 30 urn. Abbreviations: aAM,
anterior adoral membranelles; Ek, equatorial kinety; Mn, macronuclear nodule; pM, paroral
membrane; T, tail; Tk. tail kinety.

398
 CILIATES

Thigmokeronopsis crystallis Petz (1995: 138, figs 5-7)

Fig. 14.90a-c; 14.177

description: In vivo 170-300 * 60-100 um. Body elongate-ellipsoidal, dorsoventrally

c.

slightly flattened, anteriorly and posteriorly evenly rounded; buccal field large. Cirri: 26-41
frontal, in 2 rows, forming a bicorona, continuous with long midventral row of c. 41-64
cirral pairs and c. 3 single cirri posteriorly; 1 or 2 frontoterminal, or 2 buccal, 7-10 1

transverse cirri. Left postoral ventral rows of fine cirri 4-7; marginal cirral rows right and 1 1

left. Dorsal kineties 3. Adoral zone of 55-72 membranelles extending far onto right side.

Macronuclear nodules c. 110-180; micronuclei 1-6. Contractile vacuole supraequatorial,

with inconspicuous collecting canals. Movement: crawling slowly.

distribution: Weddell Sea (Petz, 1995).

Order Strombidiida

Family Strombidiidae

Laboea strobila Lohmann (1908: 171, pi. 17, fig. 14)

SYNONYMS: Conocylis helix Meunier (1910: 147, pi. 10, figs 32-35)
Strombidium slrobilum (Lohmann) Wulff (1919: 1 14, figs 21, 29, 30)

ADDITIONAL references: Kahl (1932: 499), Montagnes et al. (1988: 195), Faure-Fremiet (1924:
77)

Fig. 14.91a, b

description: In vivo 65-120 * 36-65 um [population reported by Mamaeva (1989) with

cells c. 300 um long]. Body obconical, with 4-6 distinctly notched body whorls. Equatorial
kinety extending spirally along whorls from left of oral cavity almost to posterior pole.
Ventral kinety short, on dorsal side extending from posterior end of
non-ciliferous,
equatorial kinety to antapical pole. Anterior membranelles (polykinetids) 12-18, slightly
separate from 12-22 ventral membranelles. Girdle of short extrusomes parallel to equatorial
kinety. Subpellicular platelet layer in posterior three-quarters of the body. Macronuclear
nodules 24-43 (-72); micronuclei numerous. Movement: swimming rapidly zig-zag.

distribution: Atlantic sector of the Southern Ocean (Mamaeva, 1989); Weddell Sea
(Gowing et al, 1989, as Laboea sp.); Atlantic (Faure-Fremiet, 1924; Montagnes et al.,
1988); Barents Sea, North Sea, Baltic Sea (Wulff, 1919); Pacific (Wailes, 1943).

This species can be mixotrophic due to chloroplast retention.

Spirostrombidium pseudocinctum (Wang) Petz, Song & Wilbert (1995: 123, fig. 37)

BASIONYM: Strombidium pseudocinctum Wang (1934: 62, fig. 10)

Fig. 14.106a-c

description: Fixed specimens 36-82 * 20-50 um. Body usually elongate-elliptical, with a
prominent apical protrusion in median, dorsoventrally slightly flattened. Peristomial field
rather small; oral cavity oblique; cytostome near right body margin. Equatorial kinety
curving across dorsal side and along left body margin across posterior pole to right
dorsolateral side, parallel with ventral kinety on dorsal side. Anterior adoral membranelles
26-29, inconspicuous, encircling apical protrusion, continuous with 14-17 ventral membran-
elles that curve transversely across ventral side and extend almost to right lateral margin.
Girdle of extrusomes parallel to equatorial kinety. Macronucleus 1; micronucleus 1.
Movement: swift in a spiral path.

distribution: Weddell Sea (Petz et al, 1995); Pacific (Wang, 1934).

399
 CILIATES

Spirostrombidium rhyticollare (Corliss & Snyder) Petz, Song & Wilbert (1995: 126,
figs 38, 65, 66)

BASIONYM: Strombidium rhyticollare Corliss & Snyder (1986: 43, fig. 5)

SYNONYM: Strombidium pulchrum (Leegaard) Kahl (1932: 500, fig. 81.9)

For additional synonyms, see Petz et al. (1995).

Figs 14.107a-c; 14.173

DESCRIPTION: In vivo 150-300 * 55-130 um. Body elongate-obconical. Equatorial kinety

spiralling almost twice around cell, extending from anterior left margin to posterior pole,
parallel with ventral kinety in posterior portion. Anterior adoral membranelles 18-36,
continuous with 14—40 ventral membranelles. Girdle of c. 12 um long extrusomes parallel to
equatorial kinety. Subpellicular platelet layer enclosing almost entire cell, interrupted by
somatic kineties. Macronucleus 1; micronucleus 1. Contractile vacuole possibly 1.
Movement: not fast.

distribution: Ross Sea, in the region 65.28°-76.60°S and 134.85°-165.42°W, in sea-ice and
rarely in the plankton (Petz, unpublished data); Bellingshausen Sea (Kloser, 1990); Weddell
Sea (Corliss & Snyder, 1986; Petz et al, 1995); North Sea (Leegaard, 1915); Baltic Sea
(Levander, 1894).

Strombidium acutum Leegaard (1915: 31, fig. 21)

ADDITIONAL REFERENCES: Lynn et al. (1988: 265), Montagnes et al. (1988: 195)

Fig. 14.93

description: Fixed specimens 21-53 * 18—48 um. Body obovoid. Equatorial kinety
continuous ventrally, composed of monokinetids, position supraequatorial. Ventral kinety
absent. Adoral zone of membranelles (polykinetids) prominent, 12-18 anterior membranelles
separate from 9-22 ventral membranelles. Girdle of long extrusomes in area of equatorial
kinety. Macronucleus 1.

distribution: Indian sector of the Southern Ocean (Grey et al, 1997); North Atlantic, North
Sea (Leegaard, 1915); Atlantic (Montagnes et al, 1988).

Strombidium antarcticum (Busch) Kahl (1932: 504, fig. 81.36)

BASIONYM: Buehringa antarctica Busch (1930: 84, fig. 1)

SYNONYM: Strobilidium antarcticum (Busch) Kahl sensu Hada (1970: 21), par lapsus

ADDITIONAL REFERENCE: Petz et al. (1995: 110)

Figs 14.94a-c; 14.179

description: In vivo 35-55 x 20-40 um. Body obovoid to almost spherical. Peristomial field
large; oral cavity shallow apically, separated from ventral surface by cytoplasmic wall.
Equatorial kinety continuous ventrally or with a very small gap, composed of dikinetids.
Ventral kinety usually to right of median. Adoral zone of membranelles large and forming a
closed circle in top view, 14-17 anterior membranelles continuous with 6-7 ventral
membranelles that invaginate on anterior surface. Girdle of c. 20 um long extrusomes
distinct, supraequatorial. Subpellicular platelet layer in posterior two-thirds of cell.
Macronucleus 1; micronucleus 1. Movement: usually rather slow.

distribution: Indian sector of the Southern Ocean (Hada, 1970); Ross Sea, in the region
69.98°-76.02°S and 134.85°-165.47°W, in sea-ice (Petz. unpublished data); Weddell Sea
(Petz et al, 1995); Indian Ocean (Busch, 1930).

The cytoplasm probably contains sequestered chloroplasts.

400
 CILIATES

Strombidium conicoides (Leegaard) Kahl (1932: 503, fig. 81.17)

BASIONYM: Woodania conicoides Leegaard (1915: 27, fig. 17)

Fig. 14.95

description: Fixed specimens 30-41 * 37-44 um. Subpellicular platelet layer prominent,
obconical, probably in posterior two-thirds of cell.

DISTRIBUTION: Bellingshausen Sea (Kloser, 1990); North Atlantic, North Sea (Leegaard, 1915).

a AM—/I

94a

Figures 14.93-95. Strombidium species. 14.93, S. acutum, ventral side after protargol
impregnation (from Lynn et al, 1988). 14.94, S. antarcticum, ventral side from life (a), and
ventral (b, arrow points to gap in ventral kinety, arrowhead marks ventral adoral membra-
nelles) and dorsal (c) sides after protargol impregnation (from Petz et al., 1995). 14.95, S.
conicoides, ventral side after fixation (from Kahl, 1932). Scale bars: 93 = 25 um; 94 = 20
Urn; 95 = 10 um. Abbreviations: aAM, anterior adoral membranelles; Ch, sequestered
chloroplasts; E, extrusomes (resting stage in body and extruded); Ek, equatorial kinety; P,
peristomial field; S, subpellicular platelet layer; Vk, ventral kinety.

401
 CILIATES

A poorly described species (specimens markedly deformed due to fixation) that is possibly
conspecific with S. sulcatum (q.v.).

Strombidium crassulum (Leegaard) Kahl (1932: 503, fig. 81.21)

BASIONYM: Laboea crassula Leegaard (1915: 19, fig. 9)

ADDITIONAL REFERENCE: Petz et al. (1995: 114)

Figures 14.96-100. Strombidium species. 14.96, S. crassulum, ventral side from life (a), and
ventral (b) and dorsal (c) sides after protargol impregnation (from Petz et al., 1995). 14.97,
S. diversum, ventral side after fixation (from Kahl, 1932). 14.98, S. elongatum, ventral side
after fixation(from Kahl, 1932). 14.99, 5. syowaensis, ventral side after fixation (from Hada,
1970). 14.100, S. wulffi, ventral side after protargol impregnation (from Lynn et al., 1988).
Scale bars: 96, 99 = 30 urn; 97, 98 = 20 urn; 100 = 25 urn. Abbreviations: aAM, anterior
adoral membranelles; E, extrusomes; Ek, equatorial kinety: Mn, macronuclear nodules; S,
subpellicular platelet layer; vAM, ventral adoral membranelles; Vk, ventral kinety.

402
 CILIATES

Fig. 14.96a-c

DESCRIPTION: In vivo 50-70 * 30-40 um. Body obconical, with an inconspicuous

protuberance on anterior right. Equatorial kinety discontinuous ventrally. Ventral kinety not
abutting equatorial kinety, usually distinctly left of cytostome. Anterior adoral membranelles
16-19, continuous with 13-19 ventral membranelles. Girdle of c. 17 jam long extrusomes
equatorial. Subpellicular platelet layer in posterior third to half of cell. Macronucleus 1;
micronucleus 1. Movement: moderately fast.

distribution: Ross Sea, in the region 65.28°-76.02°S and 134.85°-165.42°W, in sea-ice (Petz,
unpublished data); Weddell Sea (Petz et al., 1995); North Atlantic, North Sea (Kahl, 1932).

Strombidium diversum Busch (1930: 85, fig. 3)

SYNONYM: Strobilidium diversum (Busch) sensu Hada (1970: 27), par lapsus

ADDITIONAL REFERENCE: Kahl (1932: 504)

Fig. 14.97

description: Fixed specimens 10-39 x 10-25 um. Body globular to obconical. Subpellicular
groove (probably in the place of an equatorial kinety). Cilia
platelet layer (?) with an oblique
of adoral membranelles very long. Macronucleus 1.

distribution: Indian sector of the Southern Ocean (Hada, 1970); South Indian Ocean
(Busch, 1930).

A poorly described species.

Strombidium elongatum (Leegaard) Kahl (1932: 500, fig. 81.16)

BASIONYM: Laboea cornea Lohmann var. elongatum Leegaard (1915: 14, fig. 5d, e)

SYNONYM: Strobilidium elongatum (Leegaard) sensu Hada (1970: 28), par lapsus

Fig. 14.98

description: Fixed specimens 62-150 x 27-77 um. Body slenderly obconical, longitudinally
striped; posteriorly slightly curved, antapically pointed. Subpellicular platelet layer in
posterior five-sixths of cell. Girdle of extrusomes supraequatorial.

distribution: Indian sector of the Southern Ocean (Hada, 1970); Bellingshausen Sea
(Kloser, 1990); North Atlantic (Leegaard, 1915).

A poorly described species.

Strombidium emergens (Leegaard) Kahl (1932: 503, fig. 81.13)

BASIONYM: Laboea emergens Leegaard (1915: 21, fig. 1 1)

ADDITIONAL REFERENCE: Petz et al. (1995: 117)

Fig. 14.101a-c

description: In vivo c. 95-1 10 * 60-65 um; population of Leegaard (1915) 30-40 x 21-28
um (fixed specimens). Body elongate-ellipsoidal, without a distinct apical protrusion.
Peristomial field conspicuous; apical oral cavity separated from ventral surface by
cytoplasmic wall. Equatorial kinety continuous ventrally. Ventral kinety not abutting
equatorial kinety. Anterior adoral membranelles 14 or 15, continuous with 4 or 5 ventral
membranelles that invaginate on anterior surface and form a closed circle in top view. Girdle
of c. 12 |am long extrusomes subequatorial. Subpellicular platelet layer in posterior half of
cell. Macronucleus 1; micronucleus 1. Movement: moderately fast.

distribution: Ross Sea, in the region 71.00°-73.02°S and 135.00°-165.42°W, in sea-ice

(Petz, unpublished data); Weddell Sea (Petz et al., 1995); North Atlantic, North Sea, Baltic
Sea (Leegaard, 1915; Kahl, 1932).

403
 CILIATES

Strombidium glaciale Petz, Song & Wilbert (1995: 119, fig. 36)

Fig. 14.102a-c

DESCRIPTION: In vivo 35-45 * 30-33 um. Body obconical to ellipsoidal, with a distinct
protrusion on anterior right. Equatorial kinety usually slightly spiralling and discontinuous
ventrally. Ventral kinety extending from posterior portion almost to equatorial kinety.
Anterior adoral membranelles 16-22, continuous with 8-10 ventral membranelles. Indistinct
girdle of 6-9 urn long extrusomes usually equatorial. Subpellicular platelet layer in posterior
half of cell. Macronucleus 1; micronucleus 1. Movement: fast, spirally to and fro.

distribution: Weddell Sea (Petz et al, 1995).

Strombidium kryale Petz (1994: 186, figs 1-7)

Figs 14.103a-c; 14.180

description: In vivo 40-55 x 35-50 um. Body spherical to slightly ellipsoidal. Peristomial
field small; shallow oral cavity apical, separated from ventral surface by cytoplasmic wall.
Adoral zone inconspicuous; anterior adoral membranelles 14-17, continuous with 4-6
ventral membranelles that invaginate on anterior surface, forming a closed circle in top view.
Equatorial kinety usually continuous ventrally. Short ventral kinety in posterior third of cell.
Girdle of 8-13 um long extrusomes equatorial. Subpellicular platelet layer in posterior half
of cell. Macronucleus 1; micronucleus 1. Contractile vacuole 1. Cytoplasm containing
sequestered chloroplasts. Movement: rapid, rotating about main body axis.

distribution: Weddell Sea (Petz, 1994).

Strombidium sulcatum Claparede & Lachmann (1859: 371, pi. 13, fig. 6)

SYNONYM: Strobilidium sulcatum Claparede & Lachmann sensu Hada (1970: 21), par lapsus

ADDITIONAL REFERENCES: Borror (1963: 508), Faure-Fremiet & Ganier (1970: 207), Montagnes
et al. (1990: 319), Song et al. (2000: 333)

Fig. 14.104a-c

description: In vivo 30-60 * 18-40 um. Body obovoid to broadly ellipsoidal, with a
prominent apical protrusion in median part (disappears after fixation). Equatorial kinety
subequatorial, continuous ventrally. Ventral kinety extending from antapical pole c. halfway
to equatorial kinety. Anterior adoral membranelles 12-16, continuous with 7-9 ventral
membranelles. Girdle of prominent extrusomes (c. 15-20 um long, in bundles) parallel to
equatorial kinety. Subpellicular platelet layer in posterior half of cell. Macronucleus 1;
micronucleus 1. Contractile vacuole 1. Movement: fast.

distribution: Indian sector of the Southern Ocean (Hada, 1970); Bellingshausen Sea
(Kloser, 1990); cosmopolitan.

Strombidium syowaensis (Hada) Maeda & Carey (1985: 55, fig. 96)

BASIONYM: Strobilidium syowaensis Hada (1970: 28, fig. 40)

Fig. 14.99

description: Fixed specimens 82-160 x 67-90 um. Body obovoid, without an apical
protrusion in fixed specimens. Girdle of extrusomes supraequatorial; extrusomes grouped,
c. 20% of body length. Contractile vacuole subequatorial.

distribution: Indian sector of the Southern Ocean (Hada, 1970).

A poorly described species.

Strombidium tintinnodes Entz (1884: 387, pi. 23, figs 24-27)

SYNONYMS: Strombidium hadai Maeda & Carey (1985: 45, fig. 75)

404
 CILIATES

Strobilidium elegans Hada (1970: 28, fig. 42)

ADDITIONAL REFERENCES: Kahl (1932: 495), Alekperov & Mamaeva (1992: 10), Song &
Bradbury (1998: 769)

Fig. 14.105a-c

description: In vivo 30-54 * 15-28 urn. Body elongate-obconical, with an apical protrusion
in median part (indistinct in fixed specimens); posterior portion contractile. Equatorial kinety

101a

102a

Figures 14.101, 102. Strombidium species. 14.101, S. emergens, ventral side from life (a), and
ventral (b) and dorsal (c) sides after protargol impregnation (from Petz et al., 1995). 14.102,
S. glaciale, ventral side from life (a), and ventral (b) and dorsal (c) sides after protargol

impregnation (from Petz et al., 1995). Scale bars: 101 = 40 \im; 102 = 20 urn. Abbreviations:
Ek, equatorial kinety; Mi, micronucleus; O, oral cavity; Pr, apical protrusion; Vk, ventral
kinety.

405
 CILIATES

supraequatorial, continuous ventrally. Very short ventral kinety c. in posterior eighth of cell.
Anterior membranelles 15-18, continuous with 6 or 7 ventral membranelles.
adoral
Prominent girdle of 16-30 urn long extrusomes supraequatorial, possibly mistaken as
"longitudinal stripes" by Hada (1970). Macronucleus 1; micronucleus 1. Contractile vacuole
1. Movement: moderately to very fast.

distribution: Antarctic and/or sub- Antarctic areas of the Indian Ocean (Hada, 1970);
Pacific (Song & Bradbury, 1998); Bering Sea (Alekperov & Mamaeva, 1992); Mediterranean
Sea(Entz, 1884).

*^m8$^WWrWm Ftt{;Tfr

103a

104a

Figures 14.103, 104. Strombidium species. 14.103, S. kryale, ventral side from life (a), and
ventral (b) and dorsal (c) sides after protargol impregnation (from Petz, 1994). 14.104,
S. sulcatum, ventral side from from Faure-Fremiet & Ganier, 1970), and ventral (b)
life (a,

and dorsal (c) sides after protargol impregnation (from Song et ah, 2000). Scale bars: 103,
104 = 20 urn. Abbreviations: Ch, sequestered chloroplasts; E, extrusomes; P, peristomial
field; Pr, apical protrusion; Vk, ventral kinety.

406
 CILIATES

Strombidium wulffi Kahl (1932: 502, fig. 81.1)

BASIONYM: Strombidium striatum Wulff (1919: 113, figs 22, 27, 28), due to homonymy with Strombidium
striatum (Meunier, 1910) Kahl (1932)
SYNONYMS: Strombidium striatum Busch (1930: 86, fig. 4) sensu Kahl (1932: 504), par lapsus
Strobilidium striatum (Busch) sensu Hada (1970: 21), par lapsus

105a

106a

Figures 14.105, Strombidium and Spirostrombidium species. 14.105, Strombidium

106.
tintinnodes, ventral side from life (a), and ventral (b) and dorsal (c) sides after protargol
impregnation (from Song & Bradbury, 1998). 14.106, Spirostrombidium pseudocinctum,
ventral side from life (a, from Wang, 1934), and ventral (b) and dorsal (c) sides after
protargol impregnation (from Petz et al, 1995). Scale bars: 105a = 20 urn; 105b, c, 106a = 10
um; 106b, c = 30 urn. Abbreviations: E, girdle of extrusomes; Ek, equatorial kinety; F, fibres;
Pf, pharyngeal fibres; pM, paroral membrane; Pr, apical protrusion; Vk, ventral kinety.

407
 CILIATES

ADDITIONAL REFERENCE: Lynn et al. (1988: 264)

Fig. 14.100

description: Fixed specimens 25-90 * 18-43 um. Body obovoid, without an apical
protrusion, with several longitudinal stripes on posterior portion. Equatorial kinety distinctly
supraequatorial, continuous ventrally. Ventral kinety sigmoidal, extending from equatorial
kinety to antapical pole. Anterior adoral membranelles 11-18, continuous with 8-22 ventral
membranelles. Girdle of extrusomes parallel to equatorial kinety. Macronuclear nodules 7—
21, in 2 rings.

distribution: Indian sector of the Southern Ocean (Hada, 1970; Grey et al, 1997); cosmo-
politan.

Tontonia antarctica Petz, Song & Wilbert (1995: 130, fig. 39)

Fig. 14.92a-d

description: Trunk in vivo 55-80 x 50-75 urn, barrel-shaped; tail (caudal appendix) long,
contractile, ciliferous, detaches easily. Equatorial kinety long, extending equatorially and
across antapical pole to mid-body again, composed of dikinetids; 1 kinety on tail
(homologous to ventral kinety?). Adoral membranelles 15 or 16 anterior, separate from 22-25
ventral membranelles. Paroral membrane 1 . Subpellicular platelet layer in posterior two-thirds
of body. Extrusomes c. 13 um long, probably posterior to adoral zone and around buccal
overture. Macronuclear nodules 13-28; micronuclei several. Movement: very slow,
swimming with tail extended or rolled up and laid against body.

distribution: Ross Sea, in the region 66.00°-76.60°S and 1 62.43°- 164.97°W, in the plankton
and sometimes slush ice (Petz, unpublished data); Weddell Sea (Petz et al, 1995).

CLASS LITOSTOMATEA
Order Cyclotrichida

Family Mesodiniidae

Mesodinium pulex (Claparede & Lachmann) Stein (1867: 162)

BASIONYM: Halteria pulex Claparede & Lachmann (1859: 370, pi. 13, figs 10, 11)

ADDITIONAL REFERENCES: Kahl (1930: 127), Borror (1972: 38), Tamar (1992: 287), Foissner et
al. (1999: 139)

Fig. 14.109a-c

description: In vivo 15-35 x 10-25 um. Body pyriform; pre-equatorial furrow separating
conical anterior from globular posterior portions; blue-green to colourless. Cytostome apical,
surrounded by up to 12 bifurcated processes 23-49 short ciliary rows in area of
(retractile).
pre-equatorial furrow, bipartite. Pre-equatorial ciliary belt composed of anteriorly, laterally
and posteriorly directed bundles of long stiff and sometimes barbed cilia; equatorial kinety
belt composed of short rows of cilia which closely adjoin pellicle (difficult to observe).
Macronuclear nodules 2. Contractile vacuoles 2, supraequatorial. Terminal vacuole crescent-
shaped (function unknown). Movement: rapid backward jumps, resting with cilia stiffly
spread; attaches with oral processes to substratum.

distribution: Ross Sea, in the region 65.28°-76.02°S and 134.85°-165.47°W, in sea-ice

(Petz, unpublished data); cosmopolitan in marine and fresh waters.

408
 CILIATES

Figures 14.107-109. Spirostrombidium, Myrionecta and Mesodinium species. 14.107,

Spirostrombidium rhyticollare, ventral side from life (a), and ventral (b) and dorsal (c) sides
after protargol impregnation (from Petz et al., 1995). 14.108, Myrionecta rubra, lateral sides
from life (a, from Lindholm, 1985) and after protargol impregnation (b, from Grain et al.,
1982). 14.109, Mesodinium pulex, lateral sides from life (a, from Tamar, 1992) and after
silver impregnation (b) and detail of somatic kineties (c) (from Borror, 1972). Scale bars:
107 = 50 urn; 108a = 20 um; 108b, 109 = 10 urn. Abbreviations: aAM, anterior adoral
membranelles; Ap, apical processes; B, bristles; E, extrusomes; Eb, equatorial kinety belt;
Ek, equatorial kinety; En, endosymbiont; Mn, macronucleus; Pb, pre-equatorial kinety belt;
vAM, ventral adoral membranelles; Vk, ventral kinety.

409
 CILIATES

111a

Figures 14.110-112. Didinium, Monodinium and

Cyclotrichium species. 14.110,
D. gargantua, dorsal side from life (a) and impregnation (b) (from Petz et al.,
after protargol
1995). 14.111, M. balbianii, dorsolateral sides from life (a, from Foissner, 1979) and after
protargol impregnation (b, from Dragesco & Dragesco-Kerneis, 1986), and M. balbianii var.
nanum from life (c, lateral side, from Kahl, 1930). 14.112, C. gigas, lateral side from life
(from Faure-Fremiet, 1924). Scale bars: 110, Ilia, b = 20 urn; 112 = 100 urn; 111c = 10 urn.
Abbreviations: Ac, anterior ciliary band; Br, brosse; Cb, ciliary band; cV, contractile vacuole;
E. extrusome; Ec, equatorial ciliary band; Mn, macronucleus; Oc, oral cone; Pc, posterior
ciliary band; Sk, somatic kineties.

410
 )

CILIATES

Myrionecta rubra (Lohmann) Jankowski (1976: 168)

BASIONYM: Halteria rubra Lohmann (1908: 303, pi. 17, fig. 39)
SYNONYMS: Mesodinium rubrum (Lohmann) Hamburger & Buddenbrock (191 1: 26, non fig. 20)
Cyclotrichium meunieri Powers (1932: 75, fig. 1

ADDITIONAL REFERENCES: Kahl (1930: 127), Taylor et al. (1971: 392), Lindholm (1985: 7), Grain
et al. (1982: 8), Perriss et al. (1993; 1995: 423); Laybourne-Parry & Perriss (1995); Gibson et al.

(1997)

Figs 14.108a, b; 14.164

description: In vivo 10-70 * 10-50 um. Body barrel-shaped to obovoid, appearing to

consist of 2 hemispherical portions, with a pre-equatorial furrow separating the broader
anterior from the narrower and sometimes slightly tapering posterior portion; reddish due to
cryptophycean endosymbionts, sometimes green or colourless (same species?). Cytostome
absent; 6-8 distally bifurcated retractile processes on apex. About 30-50 bipartite kinety belt
rows; pre-equatorial belt composed of anteriorly, laterally and posteriorly directed bundles of
long and stiff cilia (bristles); equatorial belt composed of longitudinal rows of cilia which
closely adjoin pellicle. Macronuclear nodules 2; micronuclei 1 or 2. Contractile vacuole
lacking (?). Movement: jumping.

DISTRIBUTION: Indian sector of the Southern Ocean (Archer et al., 1996b; Grey et al., 1997;
Mayes et al., 1998); Ross Sea (Stoecker et al., 1993, 1995); Bellingshausen Sea (Kloser,
1990; Edwards et al, 1998); Atlantic sector of the Southern Ocean (Leakey et al., 1994;
Klaas, 1997); Weddell Sea (Corliss & Snyder, 1986; Kivi & Kuosa, 1994); also found in
brackish and saline lakes of the Vestfold Hills, East Antarctica (Perriss et al., 1993, 1995;
Laybourn-Parry et al., 1995; Gibson et al., 1997); widespread, cosmopolitan.

Probably sometimes confused with other similarly sized, reddish ciliates (e.g. Rhabdoasken-
asia; Petz, 1999).

Order Haptorida

Family Didiniidae

Cyclotrichium gigas Faure-Fremiet (1924: 37, fig. 10)

ADDITIONAL REFERENCE: Alekperov & Mamaeva (1992: 8)

Fig. 14.112

DESCRIPTION: Fixed specimens 180-265 x 205-300 urn. Body subspherical to lenticular.

Somatic kineties c. 240-250 longitudinal, not extending to apical pole; anterior portions non-
ciliferous, posterior to these 1 equatorial ciliary band of c. 35 um long cilia, with rows of
short cilia posterior to this. Macronucleus almost forming a complete ring; micronucleus 1.
Movement: rapid, rotating about main body axis.
distribution: Atlantic (Faure-Fremiet, 1924); Bering Sea (Alekperov & Mamaeva, 1992).

The species was only tentatively identified by Leakey et al. (1994) from the Atlantic sector
of the Southern Ocean. However, I found a Cyclotrichium species in sea-ice and plankton of
the Ross Sea, in the region 68.95°-76.60°S and 134.98°-162.43°W (Petz, unpublished data).

Didinium gargantua Meunier (1910: 154, pi. 15, figs 9-15)

ADDITIONAL REFERENCE: Petz et al. (1995:21)

Fig. 14.110a, b

411
 c

CILIATES

description: In vivo 70-200 * 50-120 um. Body barrel-shaped to globular, with anterior
oral cone (proboscis). Somatic kineties 50-75, longitudinal, non-ciliferous except for 2
ciliary bands (1 at anterior end of kineties and 1 in posterior quarter of body). Brosse
posterior to each ciliary band (wreath), composed of 2 or 3 short rows. Extrusomes of 3
types: rod-like cyrtocysts; fusiform pexicysts (4-6 urn long); and toxicysts 40-50 um long,
between nematodesmata. Macronucleus sausage-shaped. Contractile vacuole probably
terminal. Movement: moderately fast.

distribution: Pacific sector of the Southern Ocean (Mamaeva, 1984, 1986); Ross Sea, in the
region 68.98°-76.60°S and 134.98°-165.47°W, in sea-ice and occasionally in the plankton
(Petz, unpublished data); Atlantic sector of the Southern Ocean (Mamaeva, 1989); Weddell
Sea (Garrison & Buck, 1989a; Petz et al, 1995); Barents Sea (Meunier, 1910); Baltic Sea
(Biernacka, 1963); Pacific (Hada, 1961).

The freshwater species Didinium nasutum (?), reported by Schroder (1906a) without further
description from the Indian sector of the Southern Ocean, is considered to be conspecific
with D. gargantua.

Monodinium balbianii Fabre-Domergue (1888: 35, figs 34-50)

SYNONYM: Didinium balbianii (Fabre-Domergue) Kahl (1930: 125, fig. 18.24)

ADDITIONAL REFERENCES: Dragesco & Dragesco-Kerneis (1986: 173), Foissner et al. (1995: 235)

Fig. 14.111 a—

DESCRIPTION: In vivo 50-80 (-120) * 30-80 um. Body ovoid, with a prominent anterior oral
cone. Somatic kineties 54-80 (-100), longitudinal, non-ciliferous except for 1 ciliary band at
anterior end of kineties. Brosse posterior to ciliary band, composed of 4 or 5 short rows of
bristle-like cilia. Circumoral kinety immediately anterior to ciliary band. Extrusomes 5-10
um long, in oral cone and body. Granular mucocysts present in cortex. Macronucleus
horseshoe-shaped to reniform. Contractile vacuole in posterior end. Movement: usually very
rapidly to and fro.

distribution: Indian sector of the Southern Ocean (Hada, 1970); Weddell Sea (Garrison &
Buck, 1989a); occurs also in freshwater and soil, cosmopolitan. Records from Antarctic
freshwater lakes (Hada, 1970) and moss (Sudzuki, 1979) probably belong to a different
species (Petz & Valbonesi, unpublished data).

This ciliate is apparently highly variable and comprises several rather similar subspecies or

species. Didinium balbianii var. nanum Kahl (1930: 126, fig. 18.27) (correct identification:
Monodinium balbianii nanum), also reported from the Southern Ocean by, for example, Hada
(1970) and Kloser (1990), is only 20-35 jam long and perhaps a separate species
(Fig. 14.111c). An unidentified Monodinium sp. (Fig. 14.192) was found in an Antarctic
hypersaline lake close to the sea (Highway Lake, Vestfold Hills), at 68°29'S 78°09'E (Scott,
unpublished data).

Family Pseudotrachelocercidae

Pseudotrachelocerca trepida (Kahl) Song (1990: 161, figs 1-19)

BASIONYM: Trachelocerca trepida Kahl (1928a: 65, fig. 6a)

SYNONYM: Enchelyodon trepida (Kahl) Borror (1965: 551, figs 1, 5)

ADDITIONAL REFERENCES: Kahl (1930: 121), Wilbert & Kahan (1981: 73), Agamaliev (1983: 65),
Song (1990: 161), Petz et al. (1995: 15)

Fig. 14.113a-d

DESCRIPTION: In vivo 80-200 x 30^45 um. Body elongate, club- to spindle-shaped, anteriorly
narrowed, posteriorly rounded, contractile. Somatic kineties meridional, 19-33. Circumoral
kinety composed of a single dikinetid at anterior end of each somatic kinety. Adoral organelle

412
 CILIATES

Figures 14.113-115. Pseudotrachelocerca and Lacrymaria species. 14.113, P. trepida, lateral

side from life (a), and dorsal (b) and ventral (c) sides and oral area (d) after protargol
impregnation (from Petz et al, 1995). 14.114, L. lagenula, lateral sides from life (a) and after
protargol impregnation (b) (from Petz et al., 1995). 14.115, L. spiralis, lateral sides from life
(a) and after protargol impregnation (c, d) and cortical granulation (b) (from Petz et al,
1995). Scale bars: 113 = 20 urn; 114 = 30 urn; 115a, c = 60 um; 115d = 40 urn. Abbrev-
iations: Ao, adoral organelle; Br, brosse; Ck, circumoral kinety; cV, contractile vacuole; E,
extrusomes; Ep, excretory pore of contractile vacuole; G, cortical granules; H, head; Mi,
micronucleus; Mn, macronucleus; N, neck; PI, pharyngeal plug; Sk, somatic kinety.

413
 CILIATES

117a

Figures 14.116-118. Fuscheria, Chaenea and Loxophyllum species. 14.116, F. marina, right
side from life (a), and left (b) and right (c) sides after protargol impregnation (from Petz et
al., 1995). 14.117, C. teres, lateral side from life (a), and dorsal side (b) and detail of anterior

dorsal area (c) after protargol impregnation (from Petz et al., 1995). 14.118, L. rostratum, left
side from life (a), and right (b) and left (c) sides after protargol impregnation (from Petz et
al., 1995). Scale bars: 116a = 40 urn; 116b, c, 117a, 118b, c = 50 urn; 117b, 118a = 25 urn.

Abbreviations: Br, brosse; Ck, circumoral kinety; Cs, cytostome; cV, contractile vacuole; dK,
dorsolateral kineties; E, extrusomes; Mn, macronuclear nodule; Ne, nematodesmata; Ob, oral
bulge; pK, perioral kineties; Rs, rope-like structure; W, extrusome warts.

414
 CILIATES

(brosse) 25-50% of body length, parallel to somatic kineties (aklitolophic), composed of

irregularly arranged dikinetids (cilia c. 10 urn long), continues posteriorly as a somatic kinety
in a row of monokinetids. Extrusomes c. 4-8 urn long, fusiform, slightly bent, scattered in
body. Cortical granules colourless, in interkinetal rows. Macronucleus sausage- to narrowly
U-shaped; micronuclei few (c. 3). Contractile vacuole terminal. Movement: slow.

DISTRIBUTION: Ross Sea, in the region 65.95°-74.00°S and 134.85°-165.02°W, in sea-ice

(Petz, unpublished data); Weddell Sea (Petz et al, 1995); Atlantic (Borror, 1965); North Sea
(Kahl, 1930); Caspian Sea (Agamaliev, 1983); Red Sea (Wilbert & Kahan, 1981); Arabian
Gulf (Al-Rasheid, 2000); Pacific (Song, 1990).

Family Lacrymariidae

Lacrymaria lagenula Claparede & Lachmann (1859: 302, pi. 18, fig. 7)

See Petz et al. ( 1 995) for synonymy.

ADDITIONAL REFERENCES: Kent (1881:517), Petz et al. ( 1 995 36):

Figs 14.114a, b; 14.181

description: In vivo 60-150 * 20-30 urn. Body club-shaped, moderately contractile; neck
distinct; head and pharyngeal plug conspicuous. Somatic kineties 12-14, anteriorly
longitudinal, posteriorly spiralling; apical ends probably composed of dikinetids (brosse).
Extrusomes 4-12 um long, rod-shaped. Macronucleus ellipsoidal; micronucleus spherical.
Contractile vacuole subterminal. Movement: usually slow to moderately fast in a straight
line, rotating about main body axis.

distribution: Ross Sea, in the region 68.95°-74.03°S and 134.85°-165.02°W, in sea-ice

(Petz, unpublished data); Weddell Sea (Petz et al., 1995); North Atlantic (Claparede &
Lachmann, 1859); Mediterranean Sea (Gourret & Roeser, 1888); Pacific (Wailes, 1943).

Lacrymaria spiralis Corliss & Snyder (1986: 41, fig. 2)

ADDITIONAL REFERENCE: Petz et al. (1995: 32)

Figs 14.115a-d; 14.182

DESCRIPTION: In vivo (80-) 180-260 (-700) x (30-) 65-120 (-290) um. Body bottle- to club-
shaped, contractile, always posteriorly pointed; caudal constriction 30-40 um long;
pharyngeal plug conspicuous; broad head distinct from inconspicuous neck. Cytoplasm often
reddish brown (due to digested diatoms?), containing deep red, rarely yellow-green, food
vacuoles rendering specimens dark reddish brown to almost black, rarely orange or greenish
at low magnification. Cortical granules ruby-red (rarely almost colourless), in interkinetal
rows. Somatic kineties 14-49 on body, c. 35 ciliary rows on head. Brosse formed by 1-few
dikinetids at anterior end of each somatic kinety. Extrusomes c. 22 um long, rod-shaped.
Macronucleus long, irregularly spiralled; micronuclei 1-3 (always 3?). Contractile vacuole
subterminal. Movement: rather slow, short distances back and forth between detritus, rotates
about long axis when swimming.
DISTRIBUTION: Ross Sea, in the region 65.28°-76.02°S and 164.12°E-134.85°W, in sea-ice and
on the sea floor (Petz, unpublished data; Petz & Valbonesi, unpublished data); Weddell Sea
(Corliss & Snyder, 1986; Petz et al, 1995).

Family Acropisthiidae

Fuscheria marina Petz, Song & Wilbert (1995: 29, fig. 9)

Fig. 14.1 16a-c

description: In vivo 100-125 x 40-50 um. Body ellipsoidal; pharyngeal plug indistinct.
Somatic kineties 34-39; circumoral kinety composed of dikinetids. Brosse 2-rowed; longer
1

415
 CILIATES

row extends over c. body length (c. 38 dikinetids); second row distinctly shorter
one-third of
(c. 9 dikinetids); rows continue posteriorly as somatic kineties. Extrusomes in
both
cytostome nail-shaped, c. 1 1 urn long, those in body rod-like. Macronucleus ellipsoidal.
Contractile vacuole terminal. Movement: moderately fast.

distribution: Weddell Sea (Petz etal., 1995).

Family Trachelophyllidae

Chaenea teres (Dujardin) Kent (1881: 521, pi. 27, figs 41-44)

BASIONYM: Trachelitis teres Dujardin (1841: 400, pi. 7, fig. 14)

SYNONYM: Chaenea elongata (Claparede & Lachmann) Butschli (1889: pi. 58, fig. 7)

For additional synonyms, see Kahl (1930) and Petz et al. (1995).

ADDITIONAL REFERENCES: Morgan (1925: 629), Kahl (1930: 106), Lipscomb & Riordan (1990:
287), Petz et al. (1995:24)

Figs 14.1 17a-c; 14.183

description: In vivo 80-330 x 10-35 pm. Body elongate-cylindrical to fusiform, flexible,

slightly contractile; oral region head-like (indistinct after impregnation); oral bulge
protruding. Extrusomes 6-10 urn long, rod-shaped, in oral bulge and body. Cortical granules
colourless, between kineties. Somatic kineties 10—14, longitudinal, those on 'head' distinctly
spiralled. Circumoral kinety indistinct. Brosse 4-rowed, composed of dikinetids (cilia 10 urn
long); row 3 distinctly elongate. Macronuclear nodules > 100. Contractile vacuole terminal.
Movement: slowly gliding, rotating about main body axis when swimming in a straight line.
distribution: Ross Sea, in the region 67.98°-74.77°S and 164.12°E-134.85°W, in sea-ice
(Petz, unpublished data; Petz & Valbonesi, unpublished data); Weddell Sea (Petz et al.,
1995); Atlantic, North Sea (Hamburger & Buddenbrock, 1911); Baltic Sea (Biernacka,
1963); Mediterranean Sea (Dujardin, 1841); Caspian Sea (Agamaliev, 1983); Arabian Gulf
(Al-Rasheid, 2000).

Order Pleurostomatida

Family Litonotidae

Kentrophyllum antarcticum Petz, Song & Wilbert (1995: 50, fig. 16)

Fig. 14.1 19a-c

description: In vivo c. 200-310 * 70-100 pm. Body narrowly

broadly leaf-shaped, to
flexible, laterally distinctly flattened, with a rope-like structure along ventraland dorsal
margins, with 19-33 rigid spines around periphery of cell except in area of cytostome (in
anterior quarter of body). Somatic kineties 29-40 on right side, densely ciliated (few
anteriorly shortened). Perioral kineties 2 (1 almost surrounding body). Left side non-
ciliferous except for perioral kinety 1 and brosse (extending over entire length of body and
composed of rather widely spaced bristles). Extrusomes rod-shaped, of 2 types: short (3-5
um long), scattered in body; and long (c. 30 pm), usually around periphery. Macronuclear
nodules 2. Contractile vacuoles 3-5 in a dorsal row. Movement: slowly gliding.
distribution: Weddell Sea (Petz et al., 1995).

Litonotus antarcticus Song & Wilbert (2002: 24, figs 3, 21-23)

Fig. 14.120a-c

description: In vivo 70-150 * 30-65 pm. Body narrowly to broadly lanceolate, with c. 4

416
 CILIATES

Figures 14.119, 120. Kentrophyllum and Litonotus species. 14.119, K. antarcticum, left side
from life (a), and right (b, ciliation indicated) and left (c) sides after protargol impregnation
(from Petz et al, 1995). 14.120, L. antarcticus, left side from life (a), and right (b) and left
(c) sides after protargol impregnation (from Song & Wilbert, 2002). Scale bars: 119a = 60
um; 119b, c = 50 (j.m; 120 = 20 \im. Abbreviations: Br, brosse; cV, contractile vacuole; Mn,
macronuclear nodule; Ne, nematodesmata; Oa, oral area; pKl-3, perioral kineties 1-3; Rs,
rope-like structure; S, spine.

417
 CILIATES

Figures 14.121-123. Litonotus species. 14.121, L. emmerichi, right side from life (a), and
right (b) and left (c) sides after protargol impregnation (from Petz et ai, 1995). 14.122,
L. kopimorphus, left side from life (a), and left ventrolateral (b) and right dorsolateral (c)
sides after protargol impregnation (from Petz et ai, 1995). 14.123, L. lamella, right and left
sides from life (a, b) and after protargol impregnation (c, d) (from Foissner & O'Donoghue,
1990). Scale bars: 121a, 123a, b = 20 pm; 121b, c = 30 urn; 122 = 50 urn; 123c, d = 10 urn.
Abbreviations: Br, brosse; E, extrusome; pKl-3, perioral kineties 1-3; R, ridge; Sk, somatic
kineties.

418
 CILIATES

longitudinal grooves on left Somatic kineties 6-8 on

side, laterally flattened, contractile.
right (most anteriorly shortened) and Brosse of c. 18-20 dikinetids with
3 or 4 on left.

bristle-like cilia. Perioral kineties 3 (2 to right, 1 to left of mouth). Extrusomes rod-shaped,

4-6 um long, mainly along oral area. Macronuclear nodules (1 or ) 2; micronucleus 1.
Contractile vacuole terminal. Movement: slowly gliding.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002).

Litonotus emmerichi Petz, Song & Wilbert (1995: 43, fig. 14)

Fig. 14.121a-c

description: In vivo 60-90 * 25-30 um. Body broadly lanceolate, right side flat, left side
vaulted; oral area (slit) c. half body length; laterally flattened. Somatic kineties 5 on right,
anteriorly gradually shortened, 4 or 5 left kineties. Brosse composed of c. 10 dikinetids with
bristle-like cilia. Perioral kineties 3. Extrusomes rod-shaped, 3-5 um long. Macronuclear
nodules 2; micronucleus 1. Contractile vacuole terminal. Movement: slowly gliding.

distribution: Ross Sea, in the region 65.28°-74.70°S and 1 64. 12°E-1 34.85° W, in sea-ice (Petz,
unpublished data; Petz & Valbonesi, unpublished data); Weddell Sea (Petz et ai, 1995).

Litonotus kopimorphus Petz, Song & Wilbert (1995: 47, figs 15, 61)

Figs 14.122a-c; 14.184

description: In vivo c. 200-260 x 55-70 um. Body knife-blade-shaped; neck not distinct

from trunk; oral slit with an inconspicuous rope-like structure c. one-third of body length;
grooved by kineties). Somatic kineties 19-25 right,
laterally flattened (right side often slightly
9-15 left (anteriorly slightly shortened). Brosse of
c. 80 dikinetids in anterior half of body, in

anterior portion usually fragmented. Perioral kineties 3. Extrusomes curved-graver-shaped,

13-16 um long, along entire ventral margin (some always on antapical pole) and in body.
Cortical granules colourless, in interkinetal rows. Macronuclear nodules 2; micronucleus 1.
Contractile vacuole subterminal near ventral margin. Movement: slowly gliding.

distribution: Weddell Sea (Petz et al, 1995).

Litonotus lamella (Mtiller) Foissner, Berger, Blatterer & Kohmann (1995: 329, figs 1-8)

BASIONYM: Kolpoda lamella Miiller (1773: 56)

See Foissner et al. (1995) for synonymy.

ADDITIONAL REFERENCE: Foissner & O'Donoghue (1990: 672)

Fig. 14.123a-d

description: In vivo 50-100 (-200) * 10-25 um. Body narrowly lanceolate, posteriorly
broadly rounded; oral area c. 33-50% of body length, with a distinct ridge on left side
extending anteriorly onto neck; body laterally flattened, distinctly flexible, contractile. Right
somatic kineties 4-7, anteriorly shortened, 4 left kineties. Brosse c. one-third of body length.
Perioral kineties 3. Extrusomes rod-shaped, slightly curved, 5-7 um long, in oral area and in
posterior end. Macronuclear nodules 2; micronucleus 1. Contractile vacuole subterminal.
Movement: slowly gliding.

distribution: Indian sector of the Southern Ocean (Hada, 1970), but too superficially
described and possibly misidentified. Probably cosmopolitan in fresh water; marine records
are possibly misidentifications (Foissner et ai, 1995).

Loxophyllum rostratum Cohn (1866: 280, figs 8-1 1)

ADDITIONAL REFERENCES: Kahl (1930: 199); Petz et al. (1995: 55)

Fig. 14.1 18a-c

419
 )

CILIATES

description: In vivo 150-250 x 45-80 fim. Body narrowly to broadly lanceolate; frontal
portion bent dorsally, contractile; body laterally flattened, right side flat, left distinctly
vaulted; undernourished specimens with 2-5 inconspicuous ridges on left side. Trichocyst
warts on dorsal side, with a rope-like structure; rope-like structure also along ventral margin
and posteriorly a short distance on dorsal side in the area of the cytostome. Right somatic
kineties 11-21, densely ciliated (very few anteriorly shortened), 5-7 left kineties, 2
dorsolateral kineties. Brosse in anterior half of body, with bristle-like cilia. Perioral kineties
3, along entire ventral margin; row 2 also extends a short distance onto dorsal side.
Extrusomes (trichocysts) rod-shaped, straight to gently curved, 13-16 \im long, along ventral
margin and dorsally in warts. Macronuclear nodules 2; micronucleus 1. Contractile vacuole
subterminal. Movement: slowly gliding.

distribution: Ross Sea, in the region 66.95°-74.77°S and 134.85°-165.02°W, in sea-ice (Petz,
unpublished data); Weddell Sea (Petz et al., 1995); North Atlantic (Hamburger &
Buddenbrock, 1911); White Sea (Mereschkowsky, 1879); North Sea (Kahl, 1930); Mediterr-
anean Sea (Massuti Alzamora, 1929); Caspian Sea (Agamaliev, 1971); Arabian Gulf (Al-
Rasheid, 1997); Pacific (Song, 1993).

CLASS PHYLLOPHARYNGEA
Order Chlamydodontida

Family Kryoprorodontidae

Gymnozoum glaciate (Fenchel & Lee) Petz, Song & Wilbert (1995: 74)
BASIONYM: Spiroprorodon glacialis Fenchel & Lee (1972: 232, fig. 1)

Fig. 14.125

description: Fixed specimens 175-250 * 145-210 urn. Body ovoid to spherical. Somatic
kineties c. 38: 33 longitudinal and 5 spiral kineties equally long (no shortened kineto-
fragmon). Holotrichously ciliated, i.e. no cilia-free area between longitudinal and spiral
kineties. Nematodesmata at least 22, forming a long pharyngeal basket (cyrtos). Macronucleus
spherical. Contractile vacuole terminal.

distribution: Weddell Sea (Fenchel & Lee, 1972).

Gymnozoum sympagicum Petz, Song & Wilbert (1995: 74, fig. 22)

Figs 14.124a-d; 14.186-188

description: In vivo c. 35-65 * 25-35 um. Body asymmetrically obconical to ellipsoidal,

variable, sometimes laterally flattened;cytostome apically protruding, retractile, eccentric
and shifted somewhat to ventral side. Somatic kineties 21-25: 16-20 longitudinal, 4 long and 1

short (kinetofragmon) spiral kineties; spiral kineties equally spaced, with kinety 4 posteriorly
shortened. Conspicuous cilia-free area on right. Adoral membranelles 6-8, in a semicircle to
right of oral area. Nematodesmata 14-18, lining cytopharynx. Macronucleus 1; micronucleus
1. Contractile vacuole equatorial on left. Silverline system irregular, fine-meshed.
Movement: moderately fast, rotating about main body axis.
distribution: Ross Sea, in the region 66.00°-76.60°S and 162.43°-164.97°W, in sea-ice and
plankton (Petz, unpublished data); Weddell Sea (Petz et al., 1995).

Gymnozoum viviparum Meunier (1910: 180, pi. 20, figs 18-27; pi. 21, figs 1-24)

SYNONYMS: Spiroprorodon garrisoni Corliss & Snyder (1986: 40, fig. 1

Kryoprorodon garrisoni (Corliss & Snyder) Alekperov & Mamaeva (1992: 7)

ADDITIONAL REFERENCE: Petz et al. (1995: 70)

420
 CILIATES

Figures 14.124-126. Gymnozoum species. 14.124, G. sympagicum, right side from life (a),
right (b) and left (c) sides after protargol impregnation, and silverline system (d) after silver
nitrate impregnation (from Petz et al, 1995). 14.125, G. glaciate, right side from life (from
Fenchel & Lee, 1972). 14.126, G. viviparum, right side from life (a), and right (b) and left
(c) sides after protargol impregnation (from Petz et al., 1995). Scale bars: 124 = 20 um; 125
= 50 pm; 126 = 30 urn. Abbreviations: AM, adoral membranelles; C, cytopyge; Cs,
cytostome; cV, contractile vacuole; Kf, kinetofragmon; Lk, longitudinal somatic kineties;
Mn, macronucleus; Ne, nematodesmata; Sp, Spl, Sp4, spiral kineties 1, 4.

421
 CILIATES

Figures 14.127-129. Chlamydonella and Aegyriana species. 14.127, C. prostomata, ventral

side after protargol impregnation (a) and outline from life (b) (from Song & Wilbert, 2000a).
14.128, C. pseudochilodon, ventral (a) and dorsal (b) sides from life, and ventral (c) side
after protargol impregnation (from Petz et al., 1995). 14.129, A. paroliva, ventral sides from
life (a) and after protargol impregnation (b) (from Song & Wilbert, 2002). Scale bars: 127,

128c, 129 = 20 urn; 128a, b = 30 urn. Abbreviations: Br, brosse; Cs, cytostome; Ep,
excretory pore of contractile vacuole; pK, perioral kinety; St, stylus; Vk, ventral kineties.

422
 CILIATES

Fig. 14.126a-c

description: In vivo 65-145 * 50-105 um. Body ellipsoidal to spherical, asymmetrical,

sometimes laterally flattened; cytostome apically slightly protruding, retractile, eccentric and
shifted somewhat to ventral side. Somatic kineties 24-40: 19-35 longitudinal, 4 long and 1
very short (kinetofragmon) spiral kineties; spiral kinety 4 not shortened posteriorly, with a
distinct distance between kineties 4 and 5 (short kinetofragmon). Only a small non-ciliferous
area on rear. Adoral membranelles 8 or 9. Nematodesmata c. 14-20. Macronucleus 1;
micronucleus 1. Contractile vacuole in anterior third of body. Silverline system fine-meshed.
Movement: slow.
distribution: Ross Sea, in the region 65.95°-76.02°S and 134.85°-165.42°W, in sea-ice and
rarely in the plankton under ice (Petz, unpublished data); Weddell Sea (Corliss & Snyder,
1986; Petz et al., 1995); Barents Sea (Meunier, 1910); Caspian Sea (Agamaliev, 1976).

Family Lynchellidae

Chlamydonella prostomata Song & Wilbert (2000a: 213, figs 2, 7d-i)

Fig. 14.127a, b

description: In vivo c. 115-170 * 50-100 urn. Body usually elongate-ellipsoidal; cytostome

almost apical; a snout-like projection on anterior left; posteriorly tapering, dorsoventrally
flattened. Somatic kineties 21-29 ventral (4 preorally arched), with 1 very short right kinety
fragment; perioral kinety consisting of 5 or 6 short rows of dikinetids. Brosse dorsal on
anterior left of body. Nematodesmata 19-31. Macronucleus 1.

distribution: Weddell Sea (Song & Wilbert, 2000a).

Chlamydonella pseudochilodon (Deroux) Petz, Song & Wilbert (1995: 63, fig. 19)

BASIONYM: Chlamydonella pseudochilodon Deroux (1970: 159, figs 4-7), nomen nudum

Fig. 14.128a-c

DESCRIPTION: In vivo 40-85 x 25-40 um. Body broadly ellipsoidal, with an inconspicuous
snout-like projection on anterior dorsoventrally flattened. Somatic kineties 14 ventral (4
left;

preorally arched), with 1 very short right kinety fragment. Brosse (terminal fragment) dorsal
on anterior left end of body, composed of 8-17 kinetosomes; perioral kinety anterior of
cytostome, Y-shaped or composed of 1 long and 1 short parallel rows of monokinetids.
Nematodesmata 12-18. Macronucleus 1; micronucleus 1. Contractile vacuoles 2 (in anterior
third on right, in posterior third on left). Movement: gliding very slowly, thigmotactic.

distribution: Ross Sea, in the region 69.97°-73.00°S and 135.00°-165.02°W, in sea-ice

(Petz, unpublished data); Weddell Sea (Corliss & Snyder, 1986; Petz et al., 1995); Atlantic
(Deroux, 1970).

Order Dysteriida

Family Dysteriidae

Aegyriana paroliva Song & Wilbert (2002: 31, fig. 5)

Fig. 14.129a, b

description: In vivo 60-80 x 30-50 um. Body ellipsoidal to D-shaped. posteriorly

transversely truncate, variable; stylus (podite, attachment organelle) subterminal in middle;
oral basket (cyrtos) broad, composed of c. 26-30 nematodesmata (cytopharyngeal rods);

body dorsoventrally flattened. Somatic kineties 33-40, mainly on ventral side, the anterior

423
 CILIATES

portions of c. 7 rightmost kineties arch around the cytostome and terminate on dorsal side;

leftmost 10-15 rows very short. Perioral kineties probably in 3 fragments. Macronucleus 1.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002).

Dysteria calkinsi Kahl (1931: 258, fig. 40.28)

BASIONYM: Dysteria lanceolata Calkins (1902: 441, fig. 36), due to homonymy with Dysteria lanceolata
Claparede & Lachmann (1859)

ADDITIONAL REFERENCE: Song & Wilbert (2002: 35)

Fig. 14.130a, b

description: In vivo 30-50 * 20-25 um. Body ellipsoidal, laterally compressed; stylus
subterminal on ventral side; right side flat, right body plate anteriorly shortened and with 2
longitudinal furrows; left side vaulted. Somatic kineties: 5 in right field arched, c. 6 short
kineties in left equatorial field, with 1 very short external right-equatorial kinety. A few
short kineties near cytostome. Macronucleus 1. Movement: crawling slowly, thigmotactic.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002); Atlantic (Kahl,
1931).

Dysteria monostyla (Ehrenberg) Kahl (1931: 254, figs 42.19, 43)

BASIONYM: Euplotes monostylus Ehrenberg (1838: 380, pi. 42, fig. 14)

SYNONYM: Ervilia monostyla (Ehrenberg) Stein (1859b: 63)

ADDITIONAL REFERENCES: Faure-Fremiet (1965: 6681), Dragesco & Dragesco-Kerneis (1986:

287), Petz et al. (1995:66)

Figs 14.131a, b; 14.165

description: In vivo 80-160 x 33-65 um. Body almost rectangular, posteriorly slightly
narrowed, laterally flattened; left and right body plates of equal size, anteriorly gaping;
stylus subterminal on ventral side; cytostome in anterior 12-16% of body; oral basket
narrow. Somatic kineties: 5 in right field hook-like, c. 6-8 short kineties in left equatorial
field, 1 very short external right equatorial kinety; 6 short kineties near cytostome (1 preoral,
3 in left-frontal field, 2 circumoral). Macronucleus 1. Contractile vacuoles 2, near ventral
margin. Movement: crawling slowly.

description: Ross Sea, in the region 69.97°-76.02°S and 134.85°-165.47°W, in sea-ice

(Petz, unpublished data); Weddell Sea (Petz et al, 1995); cosmopolitan.

Order Exogenida

Family Ophryodendridae

Ophryodendron conicum Schroder (1906a: 359, pi. 27, figs 4, 5)

Fig. 14.132

description: Specimens c. 160 x 60 um. Body obconical; stalk c. 40 urn long. Tentacle-
bearing processes 4-6, each with 8-12 capitate tentacles. Macronucleus sausage-shaped.
Epizoic on copepods.

distribution: Indian sector of the Southern Ocean (Schroder, 1906a).

A poorly described species.

424
 CILIATES

131a 133

Figures 14.130-134. Dysteria, Ophryodendron, Thecacineta and Acineta species. 14.130,

D. calkinsi, right side from life (a) and after protargol impregnation (b, arrowhead marks left

equatorial kinety field) (from Song & Wilbert, 2002). 14.131, D. monostyla, left side from life
(a) and after protargol impregnation (b) (from Petz et al, 1995). 14.132, O. conicum after
fixation (after Schroder, 1906a). 14.133, T. calix, lateral side from life (from Matthes, 1956).
14.134, A. grahami after fixation (from Allgen, 1955). Scale bars: 130a, 133, 134 = 20 pm;
130b, 132 = 40 urn; 131 = 30 pm. Abbreviations: Cs, cytostome; Ek, external right equatorial
kinety; L, lorica; Pr, process; Rf, right field kineties; S, stalk; St, stylus; Te, tentacles.

425
 CILIATES

Figures 14.135-138. Conchacineta, Pelagacineta and Orthodonella species. 14.135,

C. rotunda, broad (a) and narrow (b) sides after fixation (from Allgen, 1952). 14.136,
P. campanula, top (a) and lateral sides (b) after fixation (as Tokophrya steueri, from
Schroder, 1911). 14.137, P. interrupta, top (a) and lateral sides (b) after fixation (from
Schroder, 1906a). 14.138, O. shenae, ventral side from life (a), and ventral (b) and dorsal (c)
sides after protargol impregnation (from Song & Wilbert, 2002). Scale bars: 135 = 10 (am;
136, 137 = 20 urn; 138 = 40 urn. Abbreviations: DK, dorsal kineties; Ne. nematodesmata; Sw,
swarmer; Sy, synhymenium; Ts, thecostyle.

426
 CILIATES

Family Thecacinetidae

Thecacineta calix (Schroder) Collin (1909: 1094)

BASIONYM: Acineta calix Schroder (1906a: 353, pi. 27, figs 1-3)

SYNONYMS: IThecacineta paradesmodorae Allgen (1949: 2, fig. 1)

Thecacineta subantarctica Allgen (1949: 3, fig. 3)

Thecacineta oblonga Allgen (1955: 41, fig. 3)

For additional synonyms, see Matthes (1956).

ADDITIONAL REFERENCES: Allgen (1955: 37), Matthes (1956: 477)

Fig. 14.133

description: Lorica 51-110 x 24-55 um, barrel-shaped, with 14-28 circular ridges; stalk
20-30 um long. Trophont attached to bottom of lorica. Capitate tentacles to c. 30, clustered
on apical end, to > 90 um long. Macronuclear nodules 1 or 2, spherical to ellipsoidal.
Contractile vacuole subterminal. Epizoic on nematodes. Swarmer vermiform.
distribution: Indian sector of the Southern Ocean (Schroder, 1906a); South Atlantic
(Allgen, 1949); Mediterranean Sea (Matthes, 1956).

Order Endogenida

Family Acinetidae

Acineta grahami Allgen (1955: 45, fig. 6)

Fig. 14.134

description: Fixed specimens 27-31 um diameter. Lorica almost spherical, distinctly

reticulate, lacking a stalk. Capitate tentacles very short. Epizoic on nematodes
{Neochromadora notocraspedota).
distribution: Atlantic sector of the Southern Ocean (Allgen, 1955).

A poorly described species of uncertain systematic status.

Conchacineta rotunda (Allgen) Curds (1985: 118, fig. 58)

BASIONYM: Acineta rotunda Allgen (1952: 173, figs 1-3)

SYNONYMS: Acineta ovoidea Allgen (1952: 175, figs 1, 2, 4)

Nematacineta rotunda (Allgen) Jankowski (1978: 495)

Nematacineta ovoidea (Allgen) Jankowski (1981: 86)

ADDITIONAL REFERENCE: Allgen (1955: 43)

Fig. 14.135a, b

DESCRIPTION: Fixed specimens c. 30-60 x 23-29 um. Lorica roundish, tapering posteriorly,
laterally flattened; wall transversely striated; stalk shorter than body, transversely striated.
Capitate tentacles few to at least 12, short, projecting in a single row through an opening in
the anterior two-thirds of lorica. Macronucleus spherical to ellipsoidal. Epizoic on nematodes
(Desmodora).

distribution: Atlantic sector of the Southern Ocean (Allgen, 1952); South Atlantic (Allgen,
1955).

A poorly described species.

427
 CILIATES

Pelagacineta campanula (Schroder) Jankowski (1978: 495)

BASIONYM: Tokophrya campanula Schroder (1906a: 358, pi. 27, figs 15, 16)

SYNONYMS: Tokophrya steueri Schroder (191 1: 757, figs 8, 9)

Corynophrya campanula (Schroder) Kahl (1934: 207, fig. 5.1)

For additional synonyms, see Curds (1987).

additional REFERENCES: Biernacka (1963: 65), Curds (1987: 83)

Fig. 14.136a, b

description: Specimens (11 0— ) 300-350 * (50-) 100-150 urn. Body discoidal; stalk long,
widening anteriorly to a cup-shaped lorica-like thecostyle. Capitate tentacles retractile,
encircling apical area in a continuous multi-rowed wreath. Macronucleus horseshoe-shaped
to elongate, always branched. Epizoic on copepods {Euchaeta, Metridia). Swarmer c. 90-100
x 30 um, narrowly ellipsoidal; ciliferous end tapering.

distribution: Indian sector of the Southern Ocean (Schroder, 1906a); Baltic Sea (Biernacka,
1963); Mediterranean Sea (Schroder, 1911; Fernandez-Leborans & Tato-Porto, 2000).

A poorly described species.

Pelagacineta interrupta (Schroder) Jankowski (1978: 495)

BASIONYM: Tokophrya interrupta Schroder (1906a: 355, pi. 27, figs 6-14)

SYNONYM: Corynophrya interrupta (Schroder) Kahl (1934: 207, fig. 5.2)

For additional synonyms, see Curds (1987).

additional reference: Curds (1987: 83)

Fig. 14.137a, b

description: Specimens to 300 * 100-140 um. Body discoidal; stalk long, widening
anteriorly to a cup-shaped lorica-like thecostyle. Capitate tentacles on apical area, retractile,
in 2 groups of c. 10-40. Macronucleus variable, horseshoe-, C- or X-shaped, frequently
branched; micronuclei 1 (to several?). Epizoic on copepods {Euchaeta, Metridia). Swarmer
c. 80 x 40 um, reniform; ventral side and one end ciliferous.

distribution: Indian sector of the Southern Ocean (Schroder, 1906a).

A poorly described species.

CLASS NASSOPHOREA

Order Synhymeniida

Family Orthodonellidae

Orthodonella shenae Song & Wilbert (2002: 26, figs 4, 20)

Fig. 14.138a-c

description: In vivo 120-180 x 40-70 um. Body elongate-ellipsoidal, with a snout-like

projection on anterior left; dorsoventrally flattened. Ventral kineties c. 28-45, interrupted
postorally by prominent synhymenium (composed of c. 80 dikinetids, only on ventral side);
dorsal kineties c. 19-25. Pharyngeal basket composed of c. 13 nematodesmata
(cytopharyngeal rods). Macronucleus 1. Contractile vacuoles 2, on left side. Movement:
gliding, with ventral side up and snout swinging up and down.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002).

428
 CILIATES

Figures 14.139-141. Zosterodasys, Placus and Tiarina species. 14.139, Z. kryophilus,

ventral side from life (a), and ventral (b) and dorsal (c) sides after protargol impregnation
(from Petz et al., 1995). 14.140, P. antarcticus, left side from life (a) and frontoventral side
(b) after protargol impregnation (from Petz et al., 1995). 14.141, T. antarctica, lateral side
after fixation (from Hada, 1970). Scale bars: 139, 140a = 30 nm; 140b = 50 urn; 141 = 20
p.m. Abbreviations: Ao, adoral organelle; Ck, circumoral kinety; cV, contractile vacuole;
DK, dorsal kineties; Sc, subapical cavity; Sy, synhymenium; Vk, ventral kineties.

429
 CILIATES

Figures 14.142, 143. Frontonia and Pseudocohnilembus species. 14.142, F. frigida, ventral
side from life (a), and ventral (b) and dorsal (c) sides after protargol impregnation (from
Petz et ai, 1995). 14.143, P. pusillus, ventral side from life (a, from Aescht & Foissner,
1992), and ventral (b) and dorsal (c) sides after dry silver nitrate impregnation (showing
from Foissner & Wilbert, 1981). Scale bars: 142 = 60 urn; 143 = 10 um. Abbrev-
silverlines;
iations: aM,
adoral membranelles (double membrane); C, cytopyge; Cc, caudal cilium; E,
extrusomes; Ep, excretory pore of contractile vacuole; Fv, food vacuole; Mi, micronucleus:
P, peniculi; Pk, postoral kineties; SI, silverlines; Su, suture; V, vestibular kineties.

430
 CILIATES

Zosterodasys kryophilus Petz, Song & Wilbert (1995: 59, fig. 18)

Figs 14.139a-c; 14.185

DESCRIPTION: In vivo 100-240 x 45-100, usually 140 * 55 urn. Body elongate-ellipsoidal to

obovoid, with an inconspicuous snout-like projection on anterior left; posteriorly tapering;
dorsoventrally flattened, non-contractile. Ventral kineties 25-36; dorsal kineties 21-33.
Synhymenium surrounding almost the entire cell, extending from anterior dorsal side
obliquely across ventral side to right body margin and sometimes to dorsal side again,
composed of 70-93 dikinetids. Cortical granules (mucocysts?) along kineties. Nemato-
desmata 17-26. Macronucleus ellipsoidal; micronucleus 1. Contractile vacuoles c. 6-9, along
left and posterior right body margin. Movement: slowly gliding, thigmotactic.

distribution: Weddell Sea (Petz et al., 1995).

CLASS PROSTOMATEA

Order Prorodontida

Family Placidae

Placus antarcticus Petz, Song & Wilbert (1995: 18, figs 5, 62)

Fig. 14.140a, b

description: In vivo 85-160 x 40-80 um. Body ellipsoidal to obovoid, distinctly indented in
anterior third in area of subapical cavity; cytostome apical; laterally flattened, often
deformed due to large prey. Somatic kineties 36-40, meridional to slightly spiralling. 2 C-shaped
ciliary rows in subapical cavity Adoral organelle (brosse) extending from subapical cavity to
apex (along right side of mouth). Circumoral kinety probably non-ciliferous. Extrusomes
rod-shaped, c. 14 um long, along adoral organelle and in subapical cavity. Macronucleus
usually ellipsoidal, sometimes slightly sausage-shaped; micronucleus 1. Contractile vacuole
terminal. Silverline system hexagonal, interrupted by longitudinal lines. Movement: not fast,
rotating about main body axis when swimming.

distribution: Weddell Sea (Petz et al, 1995).

Family Colepidae

Tiarina antarctica Hada (1970: 24, fig. 32)

Fig. 14.141

DESCRIPTION: Fixed specimens 45-62 x 30-37 um. Body broadly bullet-shaped, tapering at
rear, with longitudinal and oblique striae on side. Macronucleus 1.

DISTRIBUTION: Indian sector of the Southern Ocean (Hada, 1970).

A poorly described species.

431
 CILIATES

CLASS OLIGOHYMENOPHOREA

Order Peniculida

Family Frontoniidae

Frontonia frigida Petz, Song & Wilbert (1995: 79, figs 23, 58)

Figs 14.142a-c; 14.174

description: In vivo 190-300 * 55-70 um. Body usually lanceolate, dorsoventrally

flattened, appears dark low magnification and brown to green, sometimes reddish at higher
at
magnifications; pigment spot not observed. Somatic kineties 93-129, forming a distinct
suture on ventral side, which also extends a short distance onto dorsal side; 4 or 5
comparatively long vestibular kineties overlap 3 peniculi; postoral kineties 4 or 5.
Extrusomes densely spaced beneath pellicle; resting stages fusiform, in vivo c. 10 um long,
the ejected ones to 56 um long and rod-shaped. Macronucleus 1; micronuclei (1 or) 2,
reniform. Contractile vacuole in posterior 16-20% of body. Silverline system quadrangular.
Movement: slowly gliding.
distribution: Ross Sea (Petz, unpublished data); Weddell Sea (Petz et ai, 1995).

Order Philasterida

Family Pseudocohnilembidae

Pseudocohnilembus pusillus (Quennerstedt) Foissner & Wilbert (1981: 295, figs 2, 9-15)

basionym: Lembus pusillus Quennerstedt (1869: 16, fig. 6)

SYNONYMS: Pseudocohnilembus persalinus Evans & Thompson (1964: 347, figs 1-6)
Pseudocohnilembus longisetus Evans & Thompson (1964: 349, fig. 8)
Pseudocohnilembus sp. (Thompson & Croom, 1978: 54, figs 13, 14)

ADDITIONAL REFERENCES: Thompson (1965: 165), Aescht & Foissner (1992: 225), Foissner et al.

(1994:271), Song (2000: 303)

Fig. 14.143a-c

DESCRIPTION: In vivo 25-50 * 10-20 um. Body slenderly ovoid, anteriorly narrowly rounded,
posteriorly broadly rounded; oral area in anterior half of body. Somatic kineties 8-1 1; caudal
cilium elongate. Adoral membranelles 1 and 2 and paroral membrane arranged as a longit-
udinal double membrane; membranelle 3 forms a small patch of kinetosomes. Scutica
composed of c. 10 kinetids (some dikinetids), posterior to cytostome. Macronucleus 1;
micronucleus 1. Contractile vacuole slightly subterminal. Movement: moderately fast to fast,
sometimes rotating rapidly, never resting. Prefers saprobic conditions.

distribution: Atlantic sector of the Southern Ocean (Thompson & Croom, 1978); soil on
Nelly I., off Wilkes Land, East Antarctica (Thompson, 1965); cosmopolitan, also in
freshwater and saline soils. Usually found under putrid conditions, often together with
Uronema marinum.
There is considerable confusion regarding the taxonomy of the marine species of
Pseudocohnilembus. Foissner & Wilbert (1981) synonymised P. persalinus with P. pusillus.
Later, without providing a reason, Song (2000) treated P. persalinus as a separate species
and considered P. longisetus and P. marinus Thompson (1966) to be synonymous. As
important characters overlap between these 'species', they can be regarded as conspecific,
with the possible exception of P. marinus.

432
 CILIATES

145a

Figures 14.144, 145. Uronema species. 14.144, U. acutum, left side from life (a), and right
(b) and left (c) sides after protargol impregnation (from Petz et ai, 1995). 14.145, U.
antarcticum, left side from life (a), and ventral (b) and dorsal (c) sides after protargol
impregnation (from Petz et al, 1995). Scale bars: 144a, 145a = 20 urn; 144b, c, 145b, c = 10
pm. Abbreviations: AM1-3, adoral membranelles 1-3; C, cytopyge; Fp, frontal plate; Mn,
macronucleus; pM, paroral membrane; Sa, scutica.

433
 CILIATES

147a

Figures 14.146-148. Uronema and Uropedalium species. 14.146, Uronema elegans, left side
from life (a, from Maupas, 1883) and ventral side after silver nitrate impregnation (b, from
Dragesco & Dragesco-Kerneis, 1986). 14.147, Uronema marinum, left side from life (a, from
Song & Packroff, 1997) and ventral (b) and dorsal (c) sides after protargol impregnation
(from Petz et al, 1995). 14.148, Uropedalium antarcticum, ventral side after silver nitrate
impregnation (from Thompson, 1972). Scale bars: 146a = 20 urn; 146b, 147, 148 = 10 urn.
Abbreviations: AM1-3, adoral membranelles 1-3; Cs, cytostome; pM, paroral membrane; Sa,
scutica; SI, silverlines.

434
 CILIATES

Family Uronematidae

Urocyclon ovatum Song & Wilbert (2000a: 217, figs 4, 7A)

Fig. 14.150a, b

description: Fixed specimens 34-42 * 24-33 um. Body inverted-pyrifonn; frontal plate
large; cytostome subequatorial. Somatic kineties 23-28; caudal cilium elongate. Adoral
membranelle 1 reduced to 2-4 kinetosomes, distinctly separate from membranelles 2 and 3
(each composed of a small patch of kinetosomes); paroral membrane 3-shaped. Scutica
composed of 2 mono- or dikinetids. Extrusomes along somatic kineties. Macronucleus 1;
micronucleus 1. Contractile vacuole posterior.

distribution: Weddell Sea (Song & Wilbert, 2000a).

Uronema acutum Buddenbrock (1920: 344, fig. 4)

SYNONYMS: Parauronema virginianum Thompson (1967: 731, figs 1-7)

Uronema virginianum (Thompson) Foissner (1971: 47)
Uronema sp. (Fenchel & Lee, 1972: 233)

Parauronema sp. (Thompson & Croom, 1978: 53, fig. 12)

ADDITIONAL REFERENCES: Borror (1963: 495; 1965: 555), Petz et al. (1995: 88), Song & Wilbert
(2000b: 62)

Fig. 14.144a-c

DESCRIPTION: In vivo 30-70 * 13-27 um. Body fusiform, anteriorly tapering, posteriorly
narrowly rounded; frontal plate small; oral apparatus in anterior half of cell. Somatic kineties
9-14 (c. 22 kinetosomes per row); caudal cilium 13-17 um long. Adoral membranelles 3,
small; membranelle composed of 2 ciliary rows; paroral membrane indistinct. Scutica
1

consisting of c. 2 paired kinetosomes. Extrusomes in rows along somatic kineties, indistinct.

Macronuclear nodules (or 2); micronucleus 1. Contractile vacuole terminal. Movement:
1

moderately fast to fast, rotating about main body axis.

distribution: Atlantic sector of the Southern Ocean (Thompson & Croom, 1978); Weddell
Sea (Petz et al, 1995); Atlantic (Borror, 1963).

Uronema antarcticum (Thompson) Petz, Song & Wilbert (1995: 91, fig. 26)
BASIONYM: Parauronema antarcticum Thompson (1972: 273, fig. 13)

ADDITIONAL REFERENCE: Thompson & Croom (1978: 51)

Fig. 14.145a-c

description: In vivo 45-60 * 20-30 um. Body barrel-shaped to sac-like; oral apparatus in
anterior half of body, slightly indented; frontal plate distinct; laterally indistinctly flattened.
Somatic kineties 16-22 (c. 26-41 kinetosomes per row); caudal cilium 17-26 um long.
Adoral membranelles 3, small; membranelle composed of 2 ciliary rows; paroral membrane
1

inconspicuous. Scutica usually Y-shaped, composed of paired kinetosomes. Extrusomes

indistinct. Macronucleus 1; micronucleus 1. Contractile vacuole subterminal near ventral
margin. Movement: fast, resting for longer periods.

distribution: Atlantic sector of the Southern Ocean (Thompson, 1972; Thompson & Croom,
1978); Weddell Sea (Petz et al, 1995).

Uronema elegans (Maupas) Hamburger & Buddenbrock (1911, 54, fig. 60)

BASIONYM: Cryptochilum elegans Maupas (1883, 448, pi. 19, fig. 20)

ADDITIONAL REFERENCES: Czapik (1968: 225), Thompson & Kaneshiro (1968: 142), Thompson &
Croom (1978: 52), Agamaliev (1983: 89), Dragesco & Dragesco-Kerneis (1986: 347)

435
 CILIATES

Fig. 14.146a. b

DESCRIPTION: In vivo 32-90 * 20-45 um. Body reniform; oral apparatus in anterior half of
body, slightly indented; frontal plate distinct. Pellicle between kineties transversely striated.
Somatic kineties 21-27 (c. 24-30 kinetosomes per row); caudal cilium 1. Conspicuous gap
between adoral membranelle 1 (composed of 1 ciliary row) and membranelles 2 and 3.
Macronucleus 1; micronucleus 1. Contractile vacuole subterminal on ventral side.

distribution: Atlantic sector of the Southern Ocean (Thompson & Croom, 1978); Atlantic
(Czapik, 1968; Thompson & Kaneshiro, 1968); Mediterranean Sea (Maupas, 1883); Caspian
Sea (Agamaliev, 1983).

Uronema marinum Dujardin (1841: 392, pi. 7, fig. 13)

See Dragesco & Dragesco-Kerneis (1986) for synonymy.

ADDITIONAL REFERENCES: Kahl (1931: 356), Thompson (1972: 272), Coppellotti (1990: 366),
Petz etal. (1995: 95), Song & Packroff (1997: 336)

Fig. 14.147a-c

description: In vivo 20-60 x 10-40 um. Body barrel-shaped; oral area in anterior third to
half of body; frontal plate distinct, non-ciliferous. Somatic kineties 12-16; caudal cilium
elongate. Adoral membranelles 3, small; membranelle 1 composed of 1 (or 2) ciliary rows;
membranelle 2 of 2 or 3 equally long rows; paroral membrane inconspicuous. Scutica
composed of few kinetosomes. Macronucleus 1; micronucleus 1. Contractile vacuole
terminal. Movement: rapid.

distribution: Ross Sea (Coppellotti, 1990); Atlantic sector of the Southern Ocean
(Thompson, 1972; Thompson & Croom, 1978); South Atlantic (Petz et al, 1995);
cosmopolitan.

Uronema paramarinum Petz, Song & Wilbert (1995: 93, fig. 27)

Fig. 14.149a-c

description: In vivo 40-55 x 17-30 um. Body drop-shaped to pyriform, anteriorly pointed;
oral area in anterior half of body, indistinctly indented; frontal plate very small; laterally
slightly flattened. Somatic kineties 13-15 (c. 20-27 kinetosomes per row); caudal cilium
rather stiff, half as long as body. Adoral membranelles 3, small; membranelle composed of 1

1 ciliary row; membranelle 2 of 1 long and 1 very short row; paroral membrane
inconspicuous. Scutica Y-shaped, consisting of 4-6 single kinetosomes and 1 dikinetid.
Extrusomes indistinct in vivo. Macronucleus 1; micronucleus 1. Contractile vacuole terminal.
Movement: not fast, swimming in a spiral path.

distribution: Weddell Sea (Petz et al, 1995).

Uropedalium antarcticum Thompson (1972: 271, fig. 1 1)

Fig. 14.148

description: Fixed specimens 25-34 * 13-19 um. Body ellipsoidal; oral area extending over
two-thirds of body length, slightly indented in area of cytostome. Somatic kineties 12 (to 20
kinetosomes per row); caudal cilium elongate. Adoral membranelle 1 very short;
membranelle 2 composed of few widely spaced kinetosomes; membranelle 3 transversely
orientated; paroral membrane extends anteriorly only to adoral membranelle 3. Macronucleus
1. Contractile vacuole posterior.

distribution: Atlantic sector of the Southern Ocean (Thompson & Croom, 1978);
freshwater habitats, Antarctic Peninsula (Thompson, 1972; Thompson & Croom, 1978).

436
 CILIATES

149a

150a

Figures 14.149, 150. Uronema and Urocyclon species. 14.149, Uronema paramarinum, right
side from life (a), and ventral (b) and dorsal (c) sides after protargol impregnation (from
Petz et ah, 1995). 14.150, Urocyclon ovatum, ventral (a) and left (b) sides after protargol
impregnation (from Song & Wilbert, 2000a). Scale bars: 149a = 20 urn; 149b, c, 150 = 10
urn. Abbreviations: AM1-3, adoral membranelles 1-3; C, cytopyge; Cc, caudal cilium; cV,
contractile vacuole; Ep, excretory pore of contractile vacuole; Fp, frontal plate; Sa, scutica.

437
 CILIATES

151a c 152a

Figures 14.151-153. Cryptochilum and Porpostoma species. 14.151, C. reniforme, right side
from life (a), and right ventrolateral (b) and left dorsolateral (c) sides after protargol
impregnation (from Petz et al., 1995). 14.152, P. grassei, left side from life (a), and ventral
side (b) after protargol impregnation (from Petz et al., 1995). 14.153. P. notatum, right (a)
and left (b) sides from life, and ventral (c) side after protargol impregnation (from Song,
2000). Scale bars: 151a, 153a, b = 20 urn; 151b, c = 10 pm; 152 = 50 pm; 153c = 30 pm.
Abbreviations: AM1-3, adoral membranelles 1-3; Cc, caudal cilium; Mn, macronucleus; O,
oral cavity.

438
 CILIATES

Family Cryptochilidae

Cryptochilum reniforme Petz, Song & Wilbert (1995: 84, fig. 24)

Fig. 14.151a-c

DESCRIPTION: In vivo 50-55 * 25-30 um. Body reniform, anteriorly obliquely truncate,
without a frontal plate; oral cavity small, invaginated in anterior third of body; laterally
flattened. Somatic kineties 38-46, densely ciliated, curving around oral cavity on right, or 1

2 kineties on either side of oral area posteriorly shortened; caudal cilium 1, fine, rather short
(c. 16 um long). Adoral membranelles 3, small; curved paroral membrane short. Scutica
composed of c. 5-9 kinetosomes. Extrusomes (mucocysts) inconspicuous, in interkinetal
rows. Macronucleus 1; micronucleus 1. Contractile vacuole terminal. Movement: moderately
fast, rotating slightly spirally about main body axis when swimming, rests for feeding.

DISTRIBUTION: Weddell Sea (Petz et al., 1995).

Family Philasteridae

Porpostoma grassei (Corliss & Snyder) Petz, Song & Wilbert (1995: 101, fig. 30)
BASIONYM: Cohnilembus grassei Corliss & Snyder (1986: 41, fig. 3)

ADDITIONAL REFERENCE: Song & Wilbert (2000a: 219)

Fig. 14.152a, b

description: In vivo 120-350 * 40-110 um. Body slenderly bottle-shaped to fusiform,

laterally flattened, well-fed specimens roundish, often dark red (due to digested food?).
Somatic kineties 46-80; several caudal cilia slightly elongated. Adoral membranelle long, 1

slender-triangular, composed of transverse ciliary rows; membranelle 2 often inseparable

from membranelles 1 and 3; paroral membrane oblique. Scutica irregularly arranged.
Extrusomes (mucocysts) c. 4 um long. Macronucleus ellipsoidal, sometimes fragmented;
micronuclei 2. Contractile vacuole terminal. Movement: slow to moderately fast, gliding to
and fro on the substratum.

distribution: Ross Sea (Petz, unpublished data); Weddell Sea (Corliss & Snyder, 1986; Petz
et al, 1995;Song & Wilbert, 2000a).

Porpostoma notatum Mobius (1888: 90, pi. 7, figs 6, 7)

SYNONYM: Helicostoma notatum (Mobius) Kahl (1931: 367, fig. 63.10, 11)

ADDITIONAL REFERENCES: Mugard (1949: 202), Czapik & Jordan (1977: 166), Song (2000: 313,
as P. rotatum)

Fig. 14.153a-c

description: In vivo 60-200 x 20-60 um. Body elongate-fusiform, variable, tapering

anteriorly, posteriorly rounded, often slightly bent dorsally (banana-like); oral area 40-50%
of body length; cytostome conspicuous; laterally flattened. Pigment spot left of cytostome
reddish brown to black; red to orange granules scattered in cytoplasm. Somatic kineties 45-
50; 1 slightly elongate caudal cilium. Adoral membranelle 1 long, composed of 10-20
transverse ciliary portions; membranelle 2 longitudinal; membranelle 3 spoon-like; paroral
membrane bent. Scutica forming a long row of kinetosomes. Extrusomes 2-3 um long,
densely packed beneath pellicle. Macronucleus elongate, band-like; micronuclei c. 5.
Contractile vacuole terminal. Movement: very fast in a spiral path, resting for short periods.

distribution: Indian sector of the Southern Ocean (Hada, 1970); Atlantic (Mugard, 1949);
North Sea, Baltic Sea (Kahl, 1931); Mediterranean Sea (Vacelet, 1961); Pacific (Song, 2000).

439
 CILIATES

Family Orchitophryidae

Metanophrys antarctica Song & Wilbert (2002: 39, figs 8, 32-35)

Fig. 14.154a-d

DESCRIPTION: In vivo 30-45 x 20-30 nm. Body ovoid; pellicle slightly notched; oral area
c. 40% of body length. Somatic kineties 14; 1 caudal cilium c. 15 urn long. Adoral
membranelles 3, short (membranelle 1 possibly double-rowed); paroral membrane curved.
Scutica composed of few mono- and dikinetids. Extrusomes c. 2 urn long. Macronucleus 1;
micronucleus 1. Contractile vacuole terminal.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002).

Order Pleuronematida

Family Pleuronematidae

Pleuronema coronatum Kent (1881: 544, pi. 27, fig. 56)

ADDITIONAL REFERENCES: Kahl (1931: 388), Dragesco (1968: 91), Borror (1972: 58), Foissner et
al. (1994: 278), Song (2000: 316)

Fig. 14.155a-c

description: In vivo 60-100 (-150) * 30-55 um. Body elongate-ovoid to ellipsoidal, laterally
slightly flattened; oral area c. two-thirds of body length. Somatic kineties 35^48, 4-6 preoral.
Caudal cilia c. 10-15, elongate and stiff posteriorly. Adoral membranelles (oral polykinetids)
3, inconspicuous; undulating membrane prominent. Extrusomes c. 4 um long. Macronucleus
1; micronuclei 1-8. Contractile vacuole subterminal on dorsal side. Movement: moderately

fast, slightly wobbly, motionless for some seconds.

distribution: Ross Sea (Albergoni et al, 2000); Atlantic sector of the Southern Ocean
(Song & Wilbert, 2002); cosmopolitan in marine and fresh waters.

Pleuronema glaciale Corliss & Snyder (1986: 42, fig. 4)

ADDITIONAL REFERENCE: Petz et al. (1995: 97)

Fig. 14.158

description: Fixed specimens 124-168 * 42-72 um. Body elongate; oral area almost as long
as body. Somatic kineties 44-58, c. 6 preoral. Adoral membranelles 3; membranelle 1 half as
long as membranelle 2i (i.e. anterior, ±longitudinal portion of bipartite membranelle 2);
undulating membrane prominent. Macronucleus 1; micronucleus 1.

distribution: Weddell Sea (Corliss & Snyder, 1986; Petz et al., 1995); Indian sector of the
Southern Ocean (Grey et al., 1997).

Pleuronema puytoraci Groliere & Detcheva (1974: 92, figs 1-5)

ADDITIONAL REFERENCE: Petz et al. (1995: 97)

Fig. 14.156a, b

description: In vivo 90-170 * 40-60 um. Body elongate-ovoid to ellipsoidal, circular to

compressed in cross section; oral area c. 80% of body length. Somatic
laterally slightly
kineties 25-34; 2-4 preoral, postoral, with a few stiff elongate caudal cilia. Adoral
1

membranelles 3, inconspicuous (membranelle rather short); undulating membrane

1

prominent. Extrusomes indistinct in vivo. Macronucleus 1; micronuclei several. Contractile

vacuole 1, subterminal at level of cytostome. Movement: swimming moderately fast,
sometimes zig-zag, rotating about main body axis, resting for short periods.

440
 CILIATES

Figures 14.154, 155. Metanophrys and Pleuronema species. 14.154, M. antarctica, right side
(a) and transverse section of cortex (b) from life, and ventral (c) and dorsal (d) sides after
protargol impregnation (from Song & Wilbert, 2002). 14.155, P. coronatum, left side from life
(a), and left ventrolateral (b) and right dorsolateral (c) sides after protargol impregnation
(from Song, 2000). Scale bars: 154a = 10 pm; 154c, d = 20 urn; 155a = 40 pm; 155b, c = 30
pm. Abbreviations: AM1-3, adoral membranelles 1-3; Cc, caudal cilia; E, extrusome; uM,
undulating membrane.

441
 CILIATES

Figures 14.156, 157. Pleuronema and Cyclidium species. 14.156, P. puytoraci, right side
from life (a), and ventral side (b) after protargol impregnation (from Petz et al., 1995).
14.157, C. varibonneti, left side from life (a), and ventral (b) and dorsal (c) sides after
protargol impregnation (from Song, 2000). Scale bars: 156a = 40 (am; 156b = 30 urn; 157 =
15 urn. Abbreviations: AM1-3, adoral membranelles 1-3; Cc, caudal cilium; Fp, frontal plate;
pM. paroral membrane; Ps, postoral kineties; Sa, scutica.

442
 CILIATES

Figures 14.158-161. Pleuronema, Paractedoctema and Cothurnia species. 14.158,

Pleuronema glaciale, ventral side after protargol impregnation (from Corliss & Snyder, 1986).
14.159, Paractedoctema acruosa, left side from life (a), and ventral (b) and dorsal (c) sides
after protargol impregnation (from Song & Wilbert, 2000a). 14.160, Cothurnia antarctica,
(from Daday, 1913). 14.161, Cothurnia subglobosa, lateral side after
lateral side after fixation
fixation (from Daday, 1913). Scale bars: 158 = 30 urn; 159-161 = 20 urn. Abbreviations:
AM1-3, adoral membranelles 1-3; Cc, caudal cilium; Cs, cytostome; cV, contractile vacuole;
L, lorica; Mi, micronucleus; Mn, macronucleus; pM, paroral membrane; Po, preoral kineties;
S, stalk; Sa, scutica.

443
 CILIATES

169
^ 170 171

Figures 14.162-171. Uronychia, Codonellopsis, Myrionecta, Dysteria, Euplotes,

Laackmanniella, Diophrys and Cytharoides species. 14.162, Uronychia transfuga, LM ventral
side after protargol impregnation. 14.163, Codonellopsis glacialis, LM after protargol
impregnation (ptychocyliform lorica) (from Petz et al., 1995). 14.164, Myrionecta rubra, LM
from life. 14.165, Dysteria monostyla, right side LM from life. 14.166, Euplotes antarcticus,
ventral side LM from life. 14.167, Euplotes ewyhalinus, ventral side LM from life. 14.168,
Laackmanniella naviculaefera, LM loricae (b). 14.169, Diophrys appendiculata, LM ventral
side after protargol impregnation. 14.170, Cytharoides balechi, LM ventral side after protargol
impregnation. 14.171, Euplotes rariseta, LM from life, ventral side after protargol impregnation.
Scale bars: 162, 169 = 30 urn; 163, 171 = 10 Jim; 164. 167, 168 = 20 pm; 165, 166 = 40 pm; 170
= 50 pm.

444
 CILIATES

distribution: Weddell Sea (Petz et al, 1995); Black Sea (Groliere & Detcheva, 1974).

Family Cyclidiidae

Cyclidium varibonneti Song (2000: 299, figs 2, 12)

Fig. 14.157a-c

description: In vivo 15-25 10-15 um. Body barrel-shaped; ventral margin straight; frontal
*

plate conspicuous and set off; oral area c. 75% of body length. Somatic kineties 11-13,

leftmost posteriorly shortened; caudal cilium as long as body. Adoral membranelles 3, long;
paroral membrane prominent. Scutica composed of 2-4 dikinetids, in 2 distinctly separate
groups. Macronuclear nodules 2; possibly with micronucleus. Contractile vacuole
1

subterminal on ventral side. Movement: fast, jumping, motionless in-between.

distribution: Atlantic sector of the Southern Ocean (Song & Wilbert, 2002); Pacific (Song,
2000).

Paractedoctema acruosa Song & Wilbert (2000a: 215, figs 3, 7B)

Fig. 14.159a-c

description: In vivo c. 50-70 x 15-25 um. Body fusiform, anteriorly pointed, posteriorly
narrowly rounded; oral area in anterior two-thirds of body, slightly indented. Somatic
kineties 14-18; caudal cilium 1, c. half as long as body. Adoral membranelles 3, close
together; membranelle longest, slightly oblique, composed of 3 rows of kinetosomes;
1

membranelles 2 and 3 short; paroral membrane very long. Scutica composed of a short row
of kinetosomes. Macronuclear nodules 1-6; micronucleus 1. Contractile vacuole terminal.

distribution: Weddell Sea (Song & Wilbert, 2000a).

Order Sessilida

Family Vaginicolidae

Cothurnia antarctica (Daday) Warren & Paynter (1991: 27, figs 27, 28)
BASIONYM: Cothurniopsis antarctica Daday (1911: 97, fig. 1)

Fig. 14.160

description: Lorica 50-60 * 30 um, with 1-4 annular constrictions in anterior half. External
stalk 40-50 um long, with transverse folds. Endostyle and mesostyle not observed. Trophont
(zooid) extends just beyond aperture; pellicular striations inconspicuous. Macronucleus
globular. Epizoic on the ostracod Philomedes laevipes.

distribution: Bellingshausen Sea (Daday, 1913).

Cothurnia subglobosa (Daday) Warren & Paynter (1991: 49, fig. 155)
BASIONYM: Cothurniopsis subglobosa Daday (191 1: 98, fig. 2)

Fig. 14.161

description: Lorica 45-50 * 46-52 um, ovoid, smooth. External stalk 35-37 um long, with
transverse folds. Endostyle and mesostyle lacking. Trophonts 2 per lorica. Macronucleus C-
shaped. Epizoic on the ostracod Cythereis.

distribution: Bellingshausen Sea (Daday, 1913).

In addition to the species listed above, an unidentified peritrich ciliate of the Order Sessilida
was collected from the Indian sector of the Southern Ocean at 63°37'S 90°56'E (Fig. 14.193;
Scott, unpublished data).

445
 CILIATES

172 173

[t

V*
175 _ 176 *i
Figures 14.172-177. Condylostoma, Spirostrombidium, Frontonia, Holosticha and Thigmo-
keronopsis species after protargol impregnation. 14.172, C. gramriosum, LM ventral side.
14.173, S. rhyticollare, LM ventral side (from Petz et al, 1995). 14.174, F. frigida, LM
ventral side (from Petz et al., 1995). 14.75, 176, H. spindleri, LM ventral side (175) and
detail of oral area (176); arrowhead marks elongated posteriormost adoral membranelle
(from Petz et al., 1995). 14.177, T. crystallis, LM ventral side. Scale bars: 172 = 90 urn; 173,
174 = 40 pm; 175 = 10 pm; 176 = 20 pm; 177 = 30 pm.

446
 CILIATES

178 179 180

V
181 182 183 184 185

•*&
rf*

186 187 188

Figures 14.178-188. Rimostrombidium, Strombidium, Lacrymaria, Chaenea, Litonotus,

Zosterodasys and Gymnozoum species after protargol impregnation. 14.178, ./?. glacicolum,
LM ventral side. 14.179, S. antarcticum, LM ventral side. 14.180, S. kryale, LM ventral side.
14.181, L. lagenula, LM. 14.182, L. spiralis, LM. 14.183, C. feres, LM dorsal side. 14.184,
L. kopimorphus, LM right side. 14.185, Z. kryophilus, LM ventral side. 14.186-188, G.
sympagicum, LM right (186), LM right dorsolateral (187) and left (188) sides. Scale bars:
178, 181, 182, 185-188 = 20 urn; 179, 180 = 10 urn; 183, 184 = 30 urn.

447
 CILIATES

Figures 14.189-193. Codonellopsis, Cymatocylis, Monodinium and unidentified peritrich

species. 14.189, Codonellopsis gaussi, LM lorica. 14.190. Codonellopsis pusilla, SEM lorica
with agglutinated coccoliths. 14.191, Cymatocylis vanhoeffeni, LM lorica. 14.192,
Monodinium sp., SEM lateral side. 14.193, Unidentified, peritrich species (Order Sessilida),
SEM contracted specimen (lateral side). Scale bars: 189-193 = 10 urn. Abbreviations: B,
brosse; C, coccoliths; Cb, ciliary band; F, fenestra; Oc. oral cone; St, stalk.

448
 15. PROTISTA INCERTAE SEDIS
Fiona J. Scott & Harvey J. Marchant

[Class Filosea by Wolfgang Petz & Fiona J. Scott; Group Granuloreticulosa

by Patrick G. Quilty]

Hausmann & Hiilsmann (1996) discussed the development of classification schemes in proto-
zoology, including the perspicacious statement that "at present only preliminary systems can
be constructed in the field of protozoan and prot(oct)istan systematics". As with other protist
groups, the advent of molecular techniques will probably mean changes to traditional
systems (e.g. Corliss, 1994, 1998). We make no judgement on the validity of any particular
scheme, so that many Antarctic, autotrophic and heterotrophic species, genera or higher level
taxa dealt with in the following section remain of uncertain status. However, the
classification used here is that of Hausmann & Hiilsmann (1996), using Superphylum
Metakaryota and its constituent groups.

Garrison & Gowing (1993) discussed Antarctic marine protozooplankton assemblages and
provided an excellent overview of previous studies.

PHYLUM AMOEBOZOA
CLASS LOBOSEA
Order Amoebae

Amoeba spp.

A number of atestate amoebae have been observed, and while individual species are usually
difficult to identify, the accompanying images represent the range of morphologies found.
All were collected from the Southern Ocean, south of Australia, or from the coastal waters
near Davis Station, East Antarctica.

Figs 15.1a— d; 15.2a-c

CLASS FILOSEA
The Amoebozoa are a polyphyletic group comprising and testate amoebae
naked (atestate)
(testaceans, thecamoebians). However, two separate lineages whose
testate genera occur in
systematic affinities are not yet clearly resolved (e.g. Meisterfeld, 2002). Based on the shape
of their pseudopodia, i.e. filiform (distally pointed and thus called filopodia) or lobate (lobe-
like and distally ±rounded; lobopodia), they were previously assigned to Filosea or Lobosea
(Subclass Testacealobosia), respectively (e.g. Hausmann & Hiilsmann, 1996). Today, these
taxa are also considered artificial, although the morphology of the pseudopodia is an
important character in the systematics of testate amoebae (Meisterfeld, 2002). For pragmatic
reasons, however, we follow the classification of Hausmann & Hiilsmann (1996).

Filose testaceans produce an outer test (shell or lorica) made of proteinaceous material, or
siliceous scales (plates) secreted by the organisms, or formed of foreign particles
(e.g. mineral grains, detritus) embedded cement matrix (agglutinated tests). The
in a
pseudopods are retractile processes of the cell body which extend through the single
pseudostome (aperture) of the test. They are used for locomotion, shell construction, food

449
 PROTISTA INCERTAE SEDIS

Figure 15.1. Amoeba sp. indet. a, b, LM. c, d, TEM. Scale bars: a-d = 10 (am.

450
 PROTISTA INCERTAE SEDIS

Figure 15.2. Amoeba and Paulinella species, a-c, Amoeba sp. indet. a, b, TEM. c, SEM. d, P.
intermedia, SEM Test. Scale bars: a, b = 1 |im; c, d = 2 urn Abbreviations: P, pseudostome; S,
scales.

451
 PROTISTA INCERTAE SEDIS

gathering and phagocytosis, but they are usually not preserved by fixation. Other features of
the living cell include usually a single nucleus, contractile vacuoles and a zonation of the
cytoplasm in an anterior vacuolar portion with food and digestive vacuoles, a dark, band-like
pigment zone following this and a dense posterior region. However, due to the almost
complete lack of taxonomic characters of the cell body and the fact that empty tests often
predominate in samples, the shell structure is generally diagnostic for species identification.

Reproduction is by binary fission and includes the construction of a new test. Most testate
amoebae form a resting cyst and some, mostly soil- and moss-inhabiting species, produce a
short-term precyst to survive unfavorable environmental conditions.

Filose, testate amoebae are free-living and widespread in terrestrial and freshwater biotopes,
where they can occur in considerable abundance. While some species populate the interstitial
of sandy beaches, truly marine representatives are rare. Shells of filose testaceans do not
fossilise very well; often, only test fragments such as scales and spines remain, and these can
be used as indicators in palaeolimnology.

Order Euglyphida

Family Paulinellidae

Paulinella intermedia Vers (1993b: 275, fig. 1)

Fig. 15.2d

description: Test 6.5-10 * 3-6 um, elliptical; apical neck low; pseudostome 2-3 (am wide.
Test composed of transversely orientated siliceous plates, on the bowl arranged in 3-5
columns with 5-7 plates each, c. 3 plates forming the neck and few smaller plates at the
antapical end. Plates c. 1.7-3.5 * 0.7-2.0 um, rectangular, with rounded corners; surface flat.
Protoplast not observed.

distribution: Indian sector of the Southern Ocean, off Davis Station (68°34'S, 77°57'E), a
single test found on the undersurface of sea-ice (August 1997); subtropical Atlantic (Vers,
1993b).

With the exception of Gromia oviformis Dujardin, 1835 (Bowser et ah, 1996), which is no
longer regarded as a testate amoeba (Patterson et al., 2002), this is the first reliable report of
a testacean from the Southern Ocean and the second record of P. intermedia to our
knowledge.

Paulinella intermedia is possibly conspecific with Calycomonas gracilis Lohmann (1908:

291, pi. 17, fig. 13) found in the Baltic Sea. The latter differs in the shorter (3-6 * 2.5-5.5
um) and almost spherical test composed of 4 or 5 transverse rows of plates. However, as
further details of C. gracilis are lacking, we refrain from synonymising the species at present.

GROUP GRANULORETICULOSA
CLASS FORAMINIFEREA
Order Foraminiferida

Family Globorotaliidae

Neogloboquadrina pachyderma (Ehrenb.) Collen & Vella (1973: 19, pi. 1, figs 1 1, 12;
pi. 2, figs 1-3)

BASIONYM: Aristospira pachyderma Ehrenb. (Ehrenberg, 1861: 276)

SYNONYMS: Globigerina bulloides d'Orbingy var. borealis Brady (1881: 412; designated lectotype)

452
 PROTISTA INCERTAE SEDIS

Globigehna pachyderma (Ehrenb.) Brady (1884: 600, pi. 1 14, figs 19, 20)

Globorotalia pachyderma (Ehrenb.) Jenkins (1967a: 1068, fig. 2; 1967b: 195)

Globigerina borealis (Brady) Blow (1969: 316)

ADDITIONAL REFERENCES: Jenkins (1967b: 195-203), Kennett & Srinivasan (1983: 192, pi. 47,
figs 2, 6-8), Dieckmann et al. 99 1 82)
( 1 1 :

Fig. 15.3a-d

description: Organisms solitary, planktonic, c. 350

long. Test of calcite, low-
urn
trochospiral; equatorial periphery slightly periphery rounded; chambers
lobulate; axial
spherical to ovate, 4-4.5 chambers in the final whorl, increasing abruptly in size as added,
the closely embracing final chamber irregular, often a kummerform. Sutures on spiral and
umbilical sides radial, depressed; surface distinctly cancellate; thickened specimens have
surface covered with euhedral calcite crystals giving a rosette pattern; umbilicus narrow,
deep; aperture interiomarginal, umbilical to extraumbilical, a rather low arch with a thick
apertural rim.

distribution: Southern Ocean, 63.0°S 93.57°E; bipolar (Dieckmann et al, 1991); warm-
subtropical and tropical (Hillbrecht, 1996).

Neogloboquadrina pachyderma dominates foraminiferan assemblages in transitional to polar

water masses and occurs in low frequencies in warm-subtropical and tropical environments.
The species has broad tolerances for sea surface temperature and a preference for low sea
surface salinities with little seasonal change. Its preferences for dense surface waters with
low vertical temperature gradients and little stratification reflect its preferred high latitude
habitats (Hilbrecht, 1996).

This species is essentially the only planktonic foraminiferan living in Antarctic waters. The
foregoing description is a slight modification of that given by Kennett & Srinivasan (1983)
and applies mainly to thickened adult forms. Non-thickened juveniles have the same coiling
pattern, but the wall is thin and lacks the euhedral calcite rosettes so characteristic of the
adults. In polar regions, the organisms are predominantly 4-chambered, sinistrally coiled
morphotypes grading to 4.5-chambered, sinistral morphotypes in the sub-Antarctic, and to 4-
chambered, dextral morphotypes in warmer waters. It occurs in salinities as high as 82 ppm,
feed mainly on diatoms, and symbionts are not known (Hemleben et al., 1988).

GROUP ACTINOPODA
CLASS PHAEODAREA
Phaeodarians were reported from the Antarctic following the early oceanographic expeditions
(e.g.Haeckel, 1887; Peters, 1929) and a few more recent studies (e.g. Tibbs, 1969; Morley &
Stephien, 1985). Their size ranges from c. 50 jam to more than 1 cm; as a result they are often
not recovered from samples targeting small microplankton or larger zooplankton. Their
taxonomy is essentially based on siliceous skeletal morphology, and they prey on bacteria,
microalgae and protozoa. The numbers of species reported for Antarctic waters by Tibbs &
Tibbs (1986) are 125 (Pacific Antarctic), 70 (Ross Sea), 70 (Weddell Sea) and 67 (Scotia
Arc). The following phaeodarians were found to be abundant in the Weddell Sea and around
the Antarctic Peninsula (Gowing, 1989): Aulographonium anthoides Haecker, Cannosphaera
antarctica Haeckel, Challengeron swirei Murray, Coelechinus wapiticornis Haecker,
Euphysetta spp., Lirella spp., Phaeodina antarctica Cachon-Enjumet, Protocystis
micropelecus Haecker, Protocystis triangularis N.Peters and Protocystis tridens Haeckel (?
= P. acornis Haecker).

453
 PROTISTA INCERTAE SEDIS

Figure 15.3. Neogloboquadrina pachyderma. a. SEM Adult cell, ventral view. b. SEM Adult
cell, dorsal view, c, SEM Juvenile cell, ventral view, d, SEM Juvenile cell, dorsal view.
Scale bars: a-d = 100 urn.

454
 PROTISTA INCERTAE SEDIS

Figure 15.4. Actinomonas and Acanthocystis species, a, b, Actinomonas mirabilisl

Pteridomonas danica, TEM Motile cells, c, Acanthocystis paliformis, TEM Spine scale, d,
Acanthocystis perpusilla, SEM Solitary cell. Scale bars: a, b = 2 \im; c = 1 um; d = 5 \im.

455
 PROTISTA INCERTAE SEDIS

CLASS HELIOZEA
Heliozoans are cosmopolitan, free-living protozoans found in marine, brackish and fresh
waters (Febvre-Chevalier, 1985). Their basic structure comprises a spherical cell body with
fine axopodia (stiff pseudopods) radiating in all directions. Some species have
endosymbiotic algae, but most gain their nutrition solely by the ingestion of particulate
material, including other organisms. The cells of many genera are covered with overlapping
scales, and their taxonomy is based largely on EM
observation of these scales.

There have been relatively few collections of heliozoans from Antarctic waters. Takahashi
(1981) reported Pinaciophora fluviatilis Greef and an unidentified species of Acanthocystis
from the fast-ice in Ltitzow-Holm Bay, and Croome (1987) reported P. fluviatilis from a net
sample taken near Mawson Station. Croome et al. (1987) reported finding Pinaciophora
denticulata, Acanthocystis perpusilla, A. turfacea and another, unnamed Acanthocystis.

In addition to the species dealt with here, a number of other heliozoans are known from
Antarctic waters: Elaeorhanis cincta Greef (Marchant, unpublished data), Rhabdiophrys sp.
(Marchant, unpublished data) and Sticholonche spp. (Gowing & Garrison, 1991).

Order Ciliophryida

Family Ciliophrydae

Actinomonas mirabilis W.S.Kent (1880: 227)/Pteridomonas danica D.J.Patt. &

Fenchel (Patterson & Fenchel, 1985: 383, figs 1-30)

ADDITIONAL REFERENCES: Seiburth (1979: 415, pi. 23-14B), Larsen & Patterson (1990: 901),
Priesig et al. (1991: 383), Siemensma (1991: 202, fig. 17), Vers (1992a: 67, figs 32b-e, 33.1-4),
longetal. (1997: 99, fig. 5B)

Fig. 15.4a, b

description: Cells spherical to pomiform, 4-5 pm diameter, sessile or free-swimming,

uniflagellate, with tentacles in 1 or 2 rings around the flagellum. Flagellum emerging from
anterior end of cell, and a stalk c. 50 um long emerging from the posterior end. The
flagellum beats slowly, so that the cell swims with the stalk trailing behind.

distribution: Southern Ocean, south of Australia.

Vers (1992) noted that without thin-sectioning and TEM, it is virtually impossible to
distinguish Actinomonas mirabilis from Pteridomonas danica.

Order Centrohelida (Centroheliozoa)

Family Acanthocystidae

Acanthocystis paliformis Durrschmidt (1987: 59, pi. 4, figs 34-38)

ADDITIONAL reference: Vers (1992a: 22, fig. 8b)

Fig. 15.4c

description: [Only scales observed.] Spine scales 2.6 um long; plate scales 1.5 * 2.1 um.
Shaft of spine scales seated on a base 0.4 (am across; spine 0.1-0.15 pm wide, with a
truncated apex.

456
 PROTISTA INCERTAE SEDIS

distribution: Southern Ocean, south of Australia; freshwater lakes, Lake Lanalhue, Chile
(type locality) and Lake Rerewhakaaitu, New Zealand (Diirrschmidt, 1987); Baltic Sea
(Vors, 1992a).

Acanthocystis perpusilla J.B.Petersen & J.B.Hansen (1960: 550, text figs 3, 4, pi. 35, 36)

ADDITIONAL REFERENCES: Durrschmidt (1985: 69, pi. 7, figs 36, 38; 1987: 74), Croome et al.

(1987: 25, figs 7-9), Vers (1992a: 22, fig. 7f)

Fig. 15.4d

DESCRIPTION: Cells 4.8-10 pm diameter; spine scales 1.4-5.6 pm long; plate scales 1.5-2.8 *
0.7-1.5 um. Shaft of spine scales seated on a base 0.5-0.7 pm across; spine 0.07-0.11 pm
wide at centre and tapering gradually towards an apex with 2 small teeth. Plate scales mostly
ovoid, larger ones slightly constricted at their centre, patternless except for a slight axial
thickening.

distribution: Coastal waters near the Vestfold Hills, East Antarctica (Croome et al, 1987).

Acanthocystis turfacea Carter (1863: 263)

ADDITIONAL REFERENCES: Durrschmidt (1985: 58, pi. 1, figs 1-4), Croome et al. (1987: 27, figs
10-13), Siemensma (1991: 219, fig. 27), V0rs (1992a: 22, fig. 8c, d),

Fig. 15.5a

description: Cells 11-14 pm diameter; markedly bifurcate spine scales 1.8-2.7 pm long,
with base-plate 37-56 pm diameter; weakly bifurcate spine scales 4.2-11.5 pm long, with
base-plate 0.9-1.1 pm diameter. Plate scales oval, 0.9-1.5 x 6-9 pm, with slight axial
thickening.

distribution: Coastal waters off the Vestfold Hills, East Antarctica (Croome et al, 1987).

Family Heterophrydae

Heterophrys myriapoda Archer (1869: 276)

SYNONYM: Heterophrys marina Hertwig & Lesser (1874: 140)

ADDITIONAL REFERENCES: Bardele (1975: 85), Siemensma (1991: 216: fig. 25), Tobiesen (1991:
205), V0rs (1992a: 22, fig. 9a-e)

Fig. 15.5b

DESCRIPTION: Cells spherical, densely covered with pseudopodia and organic spicules
protruding through a gelatinous (spicule?) coating. Cell body (6-) 10-25 (-83) pm diameter.
Pseudopodia 10-30 pm long. Spicules needle-like, 1-13 pm long, mostly arranged radially,
the remainder forming a mat over the cell surface.

distribution: Southern Ocean, south of Australia; Norway (Tobiesen, 1991).

Under LM, sometimes appear as an ill-defined halo around the cell. Heterophrys
the spicules
is distinguished from other heliozoea, e.g. Acanthocystis, by the absence of silicious body
scales.

Order Rotosphaerida

Originally containing heliozoea that appeared to lack a centroplast and axonemes, this order
is likely to be abandoned in the future (Hutner et al., 1985). In the meantime, the following
genus is considered to be of uncertain systematic position.

457
 PROTISTA INCERTAE SEDIS

Figure 15.5. Acanthocystis, Heterophrys and Pinaciophora species, a, A. turfacea, SEM. b,

H. myriapoda, TEM. c, P. denticulata, SEM. d, P. fluviatilis, TEM Dislodged scale. Scale
bars: a-c = 5 (im; d = 2 \xm.

458
 PROTISTA INCERTAE SEDIS

Figure 15.6. Bodo, Leucocryptos, Cafeteria and Ancyromonas species, a, B. curvifilus, TEM.
b, L.marina, TEM Cell with anterior flagella. c, C. roenbergensis, TEM. d, A. sigmoides,
TEM. Scale bars: a, c = um; b, d = 2 urn.
1

459
 PROTISTA INCERTAE SEDIS

Pinaciophora denticulata H.A.Thomsen (1978b: 364, figs 5, 15, 16)

ADDITIONAL REFERENCES: Thomsen (1979: 130, figs 52, 53), Croome et al. (1987: 24, figs 3, 6)

Fig. 15.5c

description: Cell body spherical, 10-13.5 um diameter, covered with overlapping scales.
Scales oval to ±circular, 3-3.3 um diameter, with 3-8 perforations; distal surface 0.4-0.5 um
diameter. Spine scales tubular, 0.9-1.5 um long, 0.5-0.55 um diameter at apex, 0.24-0.35
um wide in mid-region, with base-plate 0.55-0.65 um diameter, attached to distal surface of
plate scales, usually peripheral to the large perforations but apparently arranged at random,
with up to 5 spine scales attached to each plate scale.

distribution: Coastal waters near the Vestfold Hills, East Antarctica (Croome, 1987); type
locality, Dybso Fjord, Denmark (Thomsen, 1978b); Peru (Gaarder et al., 1976); Japan
(Takahashi, 1981b).

Pinaciophora fluviatilis Greeff (1875: 1, figs 15, 16)

SYNONYM: Potamodiscus kalbei Gerloff (1968: 353)

ADDITIONAL REFERENCES: Gaarder et al. (1976: 346, pi. 71-76), Thomsen (1978b: 361, figs 4,
14), Takahashi (1981a: 12, pi. 7, figs 27, 28), Siemensma (1991: 273, fig. 64a-d)

Fig. 15. 5d

description: Cell body spherical, to 50 um diameter. Overlapping body scales circular to

slightly elliptical, 5.88-5.90 * 4.8-5.2 um, with 10 or 11 large perforations arranged in
concentric hexagonal rings, 0.7 um diameter, and scattered internal struts seen as black dots
under TEM.

distribution: Prydz Bay, East Antarctica; Mawson Harbour, East Antarctica; Liitzow-Holm
Bay (Takahashi, 1981a); Denmark (Thomsen, 1978); Switzerland (Penard, 1904); Gulf of
Mexico (Gaarder et al., 1976).
Thomsen (1978b) discussed important taxonomic aspects of this species, including Greeff s
(1873) short, original description and his subsequent, more detailed account (Greeff, 1875),
as well as Penard's (1904) emended description and the removal of P. fluviatilis from
Radiolaria to Heliozoea. Recently collected Antarctic samples appear to have fewer
perforations than those reported from localities in the Northern Hemisphere and summarised
in Gaarder et al. (1976).

SUPERCLASS KINETOPLASTA
Hausmann & Hiilsmann (1996) placed Subphylum Kinetoplasta in Phylum Euglenozoa.
Subsequently, a revised taxonomy (Hausmann et al., 2003) transferred Superclass
Kinetoplasta to Phylum Discicristata. In anticipation of further changes in the higher-level
systematics of the protozoa, we follow Hausmann & Hiilsmann (1996) for the two
kinetoplastid species known from Antarctic waters, but we retain them within Protista
Incertae Sedis.

The name of this superclass refers to the presence of a kinetoplast, a "segment of the unique
mitochondrion with an unusually high content of the so-called kDNA" (Hausmann &
Hiilsmann, 1996) visible by LM if material is Feulgen-stained. Cells possess 2 flagella arising
from an apical groove; one flagellum is often markedly reduced.

Rhynchomonas nasuta (A.C.Stokes) G.Klebs (Tong et al., 1997), Rhynchomonas sp.,

(Garrison & Buck, 1989b; Watanabe, 1982), Bodo saltans Ehrenb. (Tong et al., 1997) and B.
designis Skuja (Vors, unpublished data) have been collected in Antarctic waters, but they are
not dealt with in detail here due to the paucity of available material.

460
 PROTISTA INCERTAE SEDIS

CLASS BODONEA
Order Kinetoplastida

Family Bodonida

Bodo curvifilus Griessm. (Griessmann, 1913: 31, fig. 11)

ADDITIONAL REFERENCES: Vors (1992a: 43, figs 20a-i, 21.10), Patterson et al. (1993: 74, fig. 2J),
Tongetal. (1997: 99, fig. 3D)

Fig. 15.6a

DESCRIPTION: Cells oval to bean-shaped, 4-10 urn long, 2-6 }im wide, dorsoventrally
flattened, biflagellate. Flagella merging subapically, unequal; long flagellum trailing, 1-2
times cell body length; short flagellum directed forwards, 0.5-1 times cell body length.

DISTRIBUTION Antarctic Ocean, south of Australia; Disko Bay, Greenland, Baltic Sea (Vors,
1992a).

Specimens collected in the Southern Ocean were slightly smaller (3.6 x 1.8 urn) than those
previously documented.

Leucocryptos marina (Braarud) Butcher (1967: 49)

BASIONYM: Bodo marina Braarud (1935: 115, fig. 36)
SYNONYM: Chilomonas marina (Braarud) Halldal (1953: 58, fig. 21)

ADDITIONAL REFERENCES: Vers (1992a: 77, figs 37e-i, 40.1-4), Patterson et al. (1993: 82, fig. 7D)

Fig. 15.6b

DESCRIPTION: Cell asymmetrical, droplet-shaped, biflagellate. Cell body 10-30 um long, 3-7
urn diameter. Flagella subequal, inserted subapically near broadest end of cell, about as long
as the cell body. When swimming, one flagellum coils around the anterior end of the cell
body, the other trails, resulting in a spiralling motion.

distribution: Coastal waters near Davis Station, East Antarctica.

This is a common planktonic grazer (Vors 1992a).

PHYLUM CHROMISTA
SUBPHYLUM HETEROKONTA
CLASS BICOSOECIDEA
Order Bicosoecales

Family Cafeteriaceae
Diplophrys archeri J. Barker (Tong et al., 1997), Pseudobodo tremulans Griessm. and
Pseudobodo sp. (both Vors, unpublished data), have been reported from Antarctic waters, but
they are not dealt with in detail here due to the paucity of available material.

Cafeteria roenbergensis Fenchel & D.J.Patt. (Fenchel & Patterson, 1988: 19, figs 1-18)

ADDITIONAL REFERENCES: Preisig et al. (1991: 381, fig. 23. 3e, f), Vors (1992a: 55, fig. 26b, c),

Patterson et al. (1993: 79, figs 4H-J, 7A), Tongetal. (1997: 99, fig. 5G)

Fig. 15.6c

461
 PROTISTA INCERTAE SEDIS

description: Cells naked, reniform, with a slightly flattened and unornamented ventral
surface, 3-10 urn long, biflagellate, motile or sessile, uninucleate, without chloroplasts.
Flagella 5-8 urn long, inserted subapically on ventral side, often from a conspicuous pocket
over which hangs a lip-like protrusion; anterior flagellum usually directed forward, with 2
rows of fine hairs on either side; posterior flagellum trailing. Sessile cells attached to
substratum by the tip of the posterior flagellum, with the anterior flagellum beating in a
spiral fashion and directing a current of water onto the cell body. A suspension-feeding
protist.

distribution: Southern Ocean, south of Australia; Arctic Canada (Vors, 1993a); Baltic
(Fenchel & Patterson, 1988); Antarctic and Greenland (Vors, 1992a).

Cafeteria is a genus of small, bacterivorous zooflagellates that are especially common in

marine sediments and "marine snow" particles (Fenchel & Patterson, 1988). Species
concepts are based primarily on external morphology, and only the type species, C.
roenbergensis, has been characterised in detail at the ultrastructural and molecular levels.
Cafeteria cells reproduce asexually by binary division; cysts or other resting stages are
unknown, and sexual reproduction has not been described.

SUBPHYLUM HETEROKONTA
GENERA INCERTAE SEDIS
Ancyromonas sigmoides W.S.Kent (1880: 247-248)

SYNONYM: Bodo cephaloporus J.Larsen & D.J.Patt. (Larsen & Patterson, 1990: 816, figs 4h, i, 6a, b)

ADDITIONAL REFERENCES: Vors (1992a: 72, figs 34c-h, 35.5-6), Patterson et al. (1993: 74, fig.

2F, G), Tong et al. (1997, 102, figs 61, 7D)

Fig. 15. 6d

DESCRIPTION: Cells elliptical, dorsoventrally flattened, 4-5 urn long, 3-4 urn wide,
biflagellate. Flagella unequal; one flagellum inserted subapically but directed to the posterior
of the cell (i.e. trailing), the other flagellum thin, inserted anteriorly and directed forward.
A gliding heterotrophic flagellate.

distribution: Southern Ocean, south of Australia; Arctic Canada and Greenland, as well as
temperate and tropical localities (Vors 1992a).

A brief report of material cultured from a Prydz Bay marine sample included one cell of the
closely related Ancyromonas melba D.J.Patt. & A. G.B.Simpson (Tong et al, 1997).

GENERA INCERTAE SEDIS

Caecitellus parvulus (Greissm.) D.J.Patt., K.Nygaard, G.Steinb. & Turley (Vers, unpublished
data) and Ministeria maricola D.J.Patt., K.Nygaard, G.Steinb. & Turley (Tong et al., 1997)
have been collected from Antarctic waters, but they are not dealt with in detail here due to the
paucity of available material.

Amastigomonas debruynei De Saedeleer (1931: 90)

SYNONYM: Thecomonas trahens J.Larsen & D.J.Patt. (Larsen & Patterson, 1990: 924, figs 57h, 61e-f)

ADDITIONAL REFERENCES: Vers (1992a: 90, figs 34.j, 35.16), Patterson et al. (1993: 72, figs 2H, 3H)

Fig. 15.7a

description: Cells elliptical, 3-5 urn wide, 3-6 p.m long, biflagellate. Flagella inserted
apically and to one side, often difficult to see; anterior flagellum basally or completely

462
 PROTISTA INCERTAE SEDIS

enclosed by a thin organic theca that covers the cell body; posterior flagellum trailing under
the cell body.A gliding heterotrophic flagellate.
distribution: Southern Ocean, south of Australia; Arctic Canada and Greenland (Vors 1993a).

This species falls within the Apusomonadidae as proposed by Karpov & Mylnikov (1989).

Cryothecomonas armigera H.A.Thomsen, K.R.Buck, P. A. Bolt & D.L. Garrison (1991c:

1069, fig. 5)

ADDITIONAL REFERENCES: Garrison & Buck (1991: 166), Daugbjerg & Vors (1992: 35, figs 17.2-
17.4, 19.4-19.9)

Fig. 15.7b

description: Cells elliptical, dorsoventrally flattened, 12-32 um long, 7-23 urn diameter,
c. 5 um wide, biflagellate, with a characteristic anterior papilla and a longitudinal furrow
(cytosome) along one narrow side of cell. Flagella ±equal and usually equal in length to the
cell body, arising from the base of the papilla.

DISTRIBUTION: In sea-ice, near Davis Station, East Antarctica; McMurdo Sound (Stoecker et
al, 1993); type locality, Weddell Sea (Thomsen et al., 1991c).

The papilla, the anteriorly positioned nucleus with a conspicuous nucleolus and condensed
chromatin, and food vacuoles or lipid droplets are readily observed by LM (Thomsen et al.,
1991c).

Metromonas simplex (Griessm.) J.Larsen & D.J.Patt. (Larsen & Patterson, 1990: 912,
figs 57b, 58a-c)
BASIONYM: Phyllomonas simplex Griessm. (Greissmann, 1913: 33, fig. 13)

Figure 15.7. Amastigomonas and Cryothecomonas species, a, A. debruynei, TEM Cell with
anterior flagellum; posterior flagellum not visible, b, C. armigera, SEM Cell with broad
elliptical outline; furrow not visible in this view; swollen flagellar tips probably an artifact of
specimen preparation. Scale bars: a, b = 2 um.

463
 PROTISTA INCERTAE SEDIS

Figure 15.8. Metromonas and Telonema species, a, M. simplex, TEM. b. c, T. antarctia,

SEM Cells with long subequal flagella. d, T. subtile, TEM Single cell. Scale bars: a = 2 jam;
b, c = 10 |im; d = 2 jam.

464
 PROTISTA INCERTAE SEDIS

Figure 15.9. Petasaria heterolepis. a, TEM Dislodged scales, b, TEM Detail of scales; large
siliceous scales and 2 small unmineralised scales just visible in centre of image (arrows),
lying beneath large scale, c. d, SEM Collapsed cells showing large scales. Scale bars: a, c, d
= 1 urn; b = 0.5 urn.

465
 PROTISTA 1NCERTAE SEDIS

Figure 15.10. Thaumatomastix splendida. a, TEM Cell with flagella and scales, b, TEM
Scales around cell body, c, TEM Body scales, d, SEM Body scales, e, SEM Cell body. Scale
bars: a, b, e = 2 urn; c, d = 1 urn.

466
 I

PROTISTA INCERTAE SEDIS

** » A .-** ^ *««• **#'

3M A^
Figure 15.11. Thaumatomastix type A. a, SEM Single cell; 1 flagellum damaged, b, SEM
Single cell; long and short flagella visible, c, SEM Dislodged triangular body scales, d, SEM
Single cell, e, SEM Triangular, crown-like body scales. Scale bars: a, b, d = 5 urn; c, e =
1 um.

467
 PROTISTA INCERTAE SEDIS

ADDITIONAL REFERENCE: Vors (1992a: 83, figs 35.9, 35.10, 36. e), Tong et al. (1997: 103, fig. 7A)

Fig. 15.8a

DESCRIPTION: Cells solitary, ovoid, c. 3.5 fim long, c. 2.7 um diameter, uniflagellate
second flagellum is extremely short). Flagellum emerging from anterior
(if biflagellate, the
end of cell, usually crook-shaped, with the hooked distal end adhering to the substratum.
Movement gliding or nodding, with the flagellum trailing behind.
DISTRIBUTION: Southern Ocean, south of Australia; Arctic Canada and Greenland (Vors, 1993a).

Petasaria heterolepis Moestrup (1979: 86, figs 58, 59, 61-66)

ADDITIONAL REFERENCE: Nishida (1979: pi. 22, fig. 4, as "species indetermined")

Fig. 15.9a-d

description: Cells solitary, spherical, 4-6 um diameter, uniflagellate, covered by a periplast

of two types of scales. Flagellum 6-7 um long, with thin hair-like appendages (not flimmer-
hairs). Large scales siliceous, shaped like a wide-brimmed hat; basal part ±elliptical, 1.7-2.9
x 1.9-3.2 um, with a raised rim; part of the base patternless, the remainder with perforations

of various sizes; a hollow projection arises from the base near the centre, sometimes bearing
spinous ornamentation. Small scales unmineralised, ±circular, c. 0.45 um diameter, with 8
arms radiating from the centre

distribution: Southern Ocean, 58°39'S 158°47'E south of Australia; type locality,

Kaikoura, New Zealand, 42°25'S 173°42'E (Moestrup, 1979); north-western Pacific Ocean
(Nishida, 1979).

Telonema antarctica H.A.Thomsen, in Vers (1992a: 86)

additional REFERENCES: Daugbjerg& Vors (1992: 11, fig. 17.9), Brandt & Sleigh (2000: 97)

Fig. 15.8b, c

description: Cells solitary, ovoid, 10-20 um long, 4-7 um diameter, biflagellate. Flagella
equal to subequal, 1-2 times cell length, emerging from either side of an anterior
protuberance or snout, tapering distally.

distribution: Southern Ocean, south of Australia; sea-ice brine off Lucas I., East
Antarctica; southern Great Britain (Brandt & Sleigh, 2000); Arctic Canada (Daugbjerg &
Vors, 1992).

Telonema antarctica is larger than T. subtile and has longer, equal to subequal flagella.

Telonema subtile Griessm. (Griessmann, 1913: 61, fig. 23)

ADDITIONAL REFERENCES: Vers (1992a: 84, fig. 42a-l; 1993a: 121, fig. 40E), Daugbjerg & Vors
(1992: 11, fig. 17.7-17.8), Tong et al. (1997: 104, figs 6Q, 6R, 7C)

Fig. 15. 8d

description: Cells solitary, ovoid, 4-14 um long, 2-6 um diameter, biflagellate. Flagella
equal to subequal, straight, 1-2 times cell length, emerging from either side of an anterior
protuberance or snout. Cells typically swim 'backwards' with the flagella pointing away
from the direction of movement.

distribution: Coastal waters near Davis Station, East Antarctica; Southern Ocean, south of
Australia;Weddell Sea and Arctic Canada and Greenland (Vors, 1993a); Great Britain
(Brandt & Sleigh, 2000).

468
 PROTISTA INCERTAE SEDIS

Thaumatomastix fragilis H.A.Thomsen, Kosman & Ikavalko (1995: 175, figs 12-14
(holotype), 15)

description: [after Thomsen et al. (1995)] Cells solitary, ovoid, 6 um long, 4.6 urn
diameter, covered with scales, biflagellate. Flagella emerging from an anterior invagination,
dissimilar, one c. 10 urn long, naked, the other c. 6 urn long, thin, covered with minute oval
scales. Body scales uniform over the entire cell; scale base-plate triangular with rounded
corners, supporting a superstructure of branched rods that terminate in a central projection as
well as short projections along the adjoining material.

distribution: Sea-ice, Weddell Sea (Thomsen et al., 1995).

Thaumatomastix fusiformis H.A.Thomsen, Kosman & Ikavalko (1995: 175, figs 16-18;
19, 20 (holotype))
description: [after Thomsen et al. (1995)] Cells solitary, ovoid, 1 1 urn long, 8 urn diameter,
covered with scales, biflagellate. Flagella emerging from an anterior invagination, dissimilar,
one c. 12 urn long, naked, the other c. 4 um long, thin, covered with "boat-shaped" scales.
Body scales uniform over the entire cell; scale base-plate triangular with rounded corners,
supporting a simple superstructure of branched rods that fuse, resulting in small pentagonal
windows.

distribution: Sea-ice, Weddell Sea (Thomsen et al, 1995).

Thaumatomastix splendida H.A.Thomsen, Kosman & Ikavalko (1995: 175, figs 3

(holotype), 4-11)

ADDITIONAL REFERENCES: Takahashi (1981a: 11, pi. 7, fig. 30, as "sp. S"), Daugbjerg & Vors
(1992: 39, figs 18.3, 18.4, as Thaumatomastix sp. 2), Vors (1993a: fig. 23, as Thaumatomastix sp. 2)

Fig. 15.10a-e

description: Cells solitary, spherical to ovoid, slightly compressed, 6-19 um long, 6-15 um
diameter, covered with scales, biflagellate. Flagella emerging from an anterior invagination,
dissimilar, one 15-20 um long, naked, the other c. 8 um long, thin, covered with oval scales.
Body scales form a corona around the cell (LM); scale base triangular with rounded corners,
supporting 3 rods that fuse to form a conspicuous central spine c. 2 um long.

distribution: Coastal waters near Davis Station, East Antarctica; sea-ice, Weddell Sea
(Thomsen et al., 1995); Lutzow-Holm Bay (Takahashi, 1981a); Arctic Canada (Vors, 1993a).
Although Thomsen et al. splendida is especially adapted to life in sea-
(1995) stated that T.

have been observed in Southern Ocean water

ice brine channels or pockets, isolated scales
samples. Specimens from Davis Station sea-ice were usually smaller (6-10 um long) than the
holotype (12-19 um).

Thaumatomastix type A
Fig. 15.1 la-e

description: Cells solitary, ovoid, 15-22 um long, 10-1 1 um diameter, covered with scales,
biflagellate. Flagella emerging from an anterior invagination, dissimilar, one 18-20 um long,
naked, the other 8-10 um long, thin, covered with scales. Body scales form a corona around
the cell (LM); scale base triangular with rounded corners, supporting 3 rods that join in a
crown-like arrangement (Fig. 15.1 lc).

distribution: Sea-ice, Burton Lake, a coastal lagoon in the Vestfold Hills, East Antarctica.

Thomsen & Ikavalko (1997) listed 13 species of Thaumatomastix, many of which are found
in polar waters. However, the cell and scale morphology of our 'type A' appear to differ
significantly from those of recognised taxa.

469
 GLOSSARY
This glossary based largely on definitions provided in the following texts: for diatoms,
is

Medlin & Round et al. (1990: 1-58) and Hasle & Syvertsen (1997: 14-
Priddle (1990: 67-81),
20); for dinoflagellates, Steidinger & Tangen (1997: 390-416); for coccolithophores, Heimdal
(1997: 744-749); for protozoa, Corliss & Lorn (1985: 576-602), Patterson & Hedley (1992:
193-206), Hausmann & Hulsmann (1996: 301-31 1) and Throndsen (1997: 600-602).

AAC: of coccolithophores, an antapical coccolith (e.g. in Pappomonas).

aboral: relating to position, away from the oral apparatus, cf. adoral.

acicular: spindle- or needle-shaped.

acontractile: non-contractile.

acronematic: of a flagellum, terminating in a narrow hair-point.

actinopod: a ray-shaped pseudopodium.

ADBEX: (Australian) Antarctic Division Biological Experiment, R.V. Nella Dan, Southern
Ocean, south of Australia to 69°S; ADBEX I (1982), ADBEX II (1984), ADBEX III

(1985).

adhesive disc (= basal disc): of ciliates, a cup-shaped organelle at the posterior pole of a
ciliate used for attachment to a substratum.

adoral: relating to position, near the oral apparatus, cf. aboral.

adoral zone of membranelles (= oral polykinetid zone): of ciliates, a band of membran-

elles along the left and sometimes anterior sides of the oral area.

agglutinated: of ciliates, lorica wall composed entirely of foreign particles which are added
during construction and held together by an organic cement.

aklitolophic: of ciliates, type of brosse; orientation is parallel to somatic kineties and these
are not inclined upon the brosse.

ala: of prasinophytes, Pterosperma, a wing-like projection on the outer surface, pi. alae.

aloricate: without a lorica.

alveolus: of diatoms, an elongated chamber running from the central part of the valve to the
margin, open to the inside and covered by a perforated layer on the outside, pi. alveoli.

AMERIEZ: Antarctic Marine Ecosystems Research in the Ice Edge Zone, 1983, 1986 and
1988 expeditions to the Weddell and Scotia Seas (see Garrison & Buck, 1989a).

amoeba: a member of the Class Lobosea; a protist that uses pseudopodia. pi. amoebae.
amphiesmal plates (= thecal plates): of armoured dinoflagellates, plates composed of
cellulose or other polysaccharide microfibrils binding the cell.

ampulla: of ciliates and euglenoids, an anterior invagination of the cell.

anisokont: having of unequal length and/or characteristics; usually one flagellum

flagella
with flagellar hairs, the other smooth.

annulus: of diatoms, a ring of costal thickness, often surrounding one or more processes and
with a structure different from that of the rest of valve, adj. annulated.

anoxic: lacking oxygen, cf. oxic.

antapex: the posteriormost part of cell body excluding spines and similar structures.

470
 GLOSSARY

antapical plates: of dinoflagellates, thecal plates that surround and touch the antapex of the
cell.

antapical pole (= foot pole): of diatoms, mainly used for cells in stellate, radiating and
bundle-shaped colonies in which neighbouring cells are joined by valve faces of the
antapical poles, cf. apical pole.

anterior: that part of the organism which leads in forward movement.

antiligula: a small projection on diatom copulae (girdle bands) on the opposite side of the
ligula.

aperture: an opening; used in relation to loricae or tests to indicate the site of emergence of
the flagella or pseudopodia; of diatoms, the space between adjacent valves of a chain.

apex: the anteriormost part of a cell.

apical axis: the long axis of a bilateral diatom; the axis between the poles of a frustule.

apical groove: a groove at a cell apex composed of many thecal vesicles.

apical plane: perpendicular to the transapical axis.

apical plates: of dinoflagellates, those thecal plates that surround and touch the apex of the
cell.

apical pole (= head pole): of diatoms, used mainly for cells in stellate, radiating and bundle-
shaped colonies in which neighbouring cells are joined by valve faces of the antapical
poles. The apical pole is the tip of the other end of the cell. cf. antapical pole.

apical pore field: of diatoms, an area of small pores at the apices of the valves, usually in
longitudinal rows.

apical slit field: of diatoms, an area of elongate pores (slits) at the apices of the valves,
usually in longitudinal rows.

araphid: of diatoms, those with a simple raphe that runs the length of the valve.

arcuate: curved like a bow.

areola: a regularly repeated perforation through a diatom valve wall. pi. areolae.

areolate: of diatoms and dinoflagellates, ornamentation in the form of deep or shallow

depressions on thecal plates or valves.

armoured (= thecate): referring to cells with an identifiable theca or cell wall.

asexual reproduction: whereby a haploid cell produces 2 or 4 cells with the same chromosome
number as their parent.

auriculate: ear-shaped.

autotroph (-y): an organism that can manufacture its own organic requirements from
inorganic materials independent of other sources of organic substrata, adj. autotrophic;
cf. heterotroph, mixotroph.

auxotroph: a photosynthetic organism that requires only supplementary organic substances

(e.g. vitamins), adj. auxotrophic.

axial cross: of coccolithophores, a structure spanning the central area of a coccolith, with
longitudinal and transverse cross-bars, e.g. in Pappomonas weddellensis, and Papposphaera
spp.

axis: a conceptual line passing through specified parts of a cell.

axoneme: the group of microtubules (9 x 2 + 2 structure) constituting the active part of the
flagellum.

471
 GLOSSARY

axopod/axopodium: of a centrohelid heliozoan, a thin, straight pseudopodium with an

internal skeletal structure comprising microtubules.

band/segment: a single element of a diatom girdle.

basal part: of diatoms, Family Chaetoceraceae, that portion of a seta closest to the valve
face; that portion of an inner seta between its point of origin on valve face and its point
of crossing with its sibling setae.

basal siliceous layer: of diatoms, a layer that forms the basic structure of the various
components of the frustule.

BC: of coccolithophores, a body coccolith (e.g. in Pappomonas).

"Belgica": Expedition Antarctique Beige, voyage of S.Y. Belgica, 1897-99, western

Antarctic Peninsula.

benthic: referring to organisms living on the sea-bed.

biarcuate: having two adjoining bow-like curves.

bifurcated: forked, divided into two parts.

bilabiate process: of diatoms, a process consisting of an shorter or longer external tube,

sometimes reduced to a low ring (LM), and an internal part with a longer or shorter stalk
and a trapezoidal end-piece closed at the tip but open at each of the two slanting sides by
a longitudinal split.

bilateral/bilateral symmetry: of diatoms, having valve striae arranged basically in relation

to a line, so that the cell appears to have identical left and right halves.

binary fission: an asexual mode of reproduction.

biotope: the living space of a community of organisms (plants, animals and microorganisms).

blue-green algae: members of the Class Cyanophyceae.

bloom: an overabundant growth of organisms, often caused by nutrient enrichment.

box scale: of prasinophytes, a box-shaped, organic scale on the outer surface of the cell body.

BRATEGG: Norwegian Antarctic Expedition, 1947-48.

bristles: of ciliates, stiff and rather immobile cilia (long or short) of several kinds and
functions.

broad girdle view: a diatom frustule seen from the broad side.

brosse (= dorsal brush, terminal fragment): of ciliates, one or several, often short kineties
near anterior of cell (usually dorsal), often orientated obliquely to the long body axis.

buccal apparatus: of ciliates, see oral apparatus.

buccal cirrus: of spirotrich ciliates, usually one or a few cirri close to the right and often
near the anterior end of the paroral membrane.

c: circa (about).

calyx: of coccolithophores, a flaring structure at the tip of a process, e.g. in Papposphaera

spp.

campanulate: bell-shaped.

canal raphe: of diatoms, a system consisting of raphe canal, fibula and raphe keel.

cancellate: of Granuloreticulosa, latticed.

472
 GLOSSARY

caneolith: of coccolithophores, a disc- or bowl-shaped heterococcolith with a lath-filled

central area (e.g. in Syracosphaera).

capitate: having a knob-like head or tip.

caudal: ofciliates, relating to or in the position of a tail.

caudal cilium: ofciliates, a distinctly longer somatic cilium at or near the posterior pole of a
cell body.

caudal cirri: of spirotrich ciliates, at or near the posterior margin of dorsal surface and at

the ends of dorsal kineties.

cell: a mass of cytoplasm bound by plasma membrane; see also prokaryote, eukaryote.

central nodule: of diatoms, Family Bacillariaceae, a bridge of silica separating two raphe
slits, often thicker than the rest of the valve.

central pore: of diatoms, Family Bacillariaceae, a pore-like expansion of the central raphe
ending.

central raphe ending: of diatoms, Family Bacillariaceae, the central end of a raphe slit,

when the raphe consists of two slits.

centric: referring to diatoms of the Order Centrales.

centrohelid: a type of heliozoan with thin, parallel-sided arms with prominent extrusomes.

centroplast: of a centrohelid heliozoan, the single, centrally located, microtubule organising

centre (MTOC) from which all axopods arise.

CFC: of coccolithophores, a circum-flagellar coccolith (e.g. in Pappomonas).

chains: of diatoms, cells joined by threads from strutted processes (Thalassiosira), by

external parts of strutted processes (Skeletonema), by abutting of valve faces
(Leptocylindrus), by external part of the single labiate process, contiguous area and
claspers (Rhizosolenia), by bipolar elevations (Eucampia), by bipolar elevations with
spines (Ceratulina), by setae (Chaetoceros) or by a marginal ridge (Lithodesmium).

chlorophyll: a group of green pigments having the vital function of absorbing light energy
for photosynthesis.

chlorophyte (= green alga): a member of the Class Chlorophyceae.

chloroplast: a membrane-bound cell organelle containing chlorophyll.

choanoflagellate: a member of the Class Craspedophyceae/Choanoflagellida.

chromatin: chromosomal material consisting of DNA and proteins within the cell nucleus
which becomes deeply stained with basic dyes.

chromosome: an assemblage of molecular DNA in the nuclei of eukaryotic cells, containing

genetic information.

chrysophyte (= golden-brown flagellate): a member of the Class Chrysophyceae.

ciliate: a member of the Phylum Ciliophora.

ciliferous: ofciliates, bearing cilia.

cilium: of ciliates, a hair-like projection associated with locomotion, having a 9 x 2 + 2

axonemal arrangement as found in flagella. pi. cilia.

ciliary row: see kinety.

ciliature: the total assembly of oral and somatic cilia or cirri; may be arranged in patches,
kineties, membranes or membranelles.

473
 GLOSSARY

cingulum (= girdle, transverse groove): of dinoflagellates, a furrow encircling a

dinoflagellate cell once or several times, either circular, displaced, median, postmedian
or premedian in location; of diatoms, a portion of a diatom girdle associated with a single
valve; part of the frustule between upper and lower valves, adj. cingular.

circular cingulum: of dinoflagellates, a cingulum that is not displaced and in which the
proximal end meets the distal end.

circumoral: of ciliates, surrounding the oral area.

cirrus: of ciliates, compound somatic ciliature; a cluster of cilia functioning as a single unit
in locomotion or food collection, but with no common enveloping membrane, pi. cirri.

claspers: of diatoms, Family Rhizosoleniaceae, a pair of membranous structures that unite

adjacent cells by wrapping around the otaria.

clavate: club-shaped.

coccoid: a cell with a firm outer wall, without flagella.

coccolith: plate-like, calcified scales formed in vesicles of the endoplasmic reticulum of

coccolithophorid haptophytes.

coccolithophore: a member of Order Coccolithophorales.

coccosphere: the spherical cell of a coccolithophore.

collar: a hollow, cone-shaped circle of minute, finger-like processes projecting from the
apex of choanoflagellate cells.

colony: a localised population of organisms derived from a single cell.

connecting band (= pleura): of diatoms, an element in the middle of the girdle when
intercalary bands are present, or any element when intercalary bands are absent.

contiguous area: of diatoms, Family Rhizosoleniaceae, part of the ventral side of the valve
contiguous with the adjacent valve of linked cells, usually delimited by low, marginal
ridges.

contractile vacuole: a membrane-bound cavity located peripherally in the cell, associated

with the expulsion of fluid and serving as an osmoregulator in the cytoplasm.

copula: see intercalary band, girdle band.

cortex: of ciliates, the complex outer cell layer including pellicle and infraciliature.

cortical granules: of ciliates, small, often refractile granules in the periphery of a cell,
sometimes corresponding to extrusomes (e.g. mucocysts). Colour, shape and arrangement
are important species characteristics.

costa: of diatoms, a siliceous rib on valve surface; of choanoflagellates, siliceous strips or

rods from which the lorica is built.

coxlielliform: of tintinnid ciliates, lorica shape as in genus Coxliella.

crenulate: with a regularly indented margin.

cribrum: of diatoms, velum of a valve perforated by regularly arranged pores, pi. cribra.

cruciform: cross-shaped.

cristallite: of coccolithophores, a minute crystal.

cryptophyte: a member of the Class Cryptophyceae.

cuneate: wedge-shaped.

cyanophyte (= blue-green alga, cyanobacterium): a member of the Class Cyanophyceae.

cyrtocyst: of ciliates, a type of subpellicular extrusome found in Didinium.

474
 GLOSSARY

cyrtos: of ciliates, a type of tubular cytopharyngeal apparatus; often curved, with walls
strengthened by longitudinally arranged nematodesmata. Derived from apically located
kinetosomes and lines with extensions of postciliary microtubules; does not contain
toxicysts.

cytopharynx: of ciliates, a non-ciliated, tubular passageway leading from the cytostome into
the inner cytoplasm of the organism, adj. cytopharyngeal.

cytoplasm: that part of the living contents of a cell not located within the nucleus,
adj. cytoplasmic.

cytostome: of ciliates, a cell mouth or oral opening; a two-dimensional aperture, most

commonly open, through which food passes into the cytoplasm.

cyst: a bladder or bag-like structure that may contain the resting stage of an organism.

decussate: in alternating pairs.

detritus: fragments of dead biological material before, during and after decay, sometimes
incorporating inorganic matter.

diatom: a member of the Class Diatomophyceae/Bacillariophyceae.

dikinetid: of ciliates, a morphological complex of two neighbouring (paired) basal bodies

(kinetosomes) along with associated cilia and infraciliary organelles.

dinoflagellate: a member of the Class Dinophyceae.

diploid: a cell (zygote) that has a nucleus with two sets of chromosomes, abbrev. 2n.

dithecate: of coccolithophores, cells having two discrete layers of different types of

coccoliths, the endothecal and exothecal layers.

dorsal: the uppermost side, or back, of an organism.

dorsal brush: of ciliates, one or several short kineties near the anterior end of a cell.

double-eurystomous type: of euplotid ciliates, a type of dorsal silverline system; one

longitudinal silverline in the middle between each of two dorsal kineties connected with
transverse silverlines so that two longitudinal rows of equally long rectangles are formed.

double-patella type: of euplotid ciliates, a type of dorsal silverline system; one longitudinal
silverline eccentric between each of two dorsal kineties connected with transverse
silverlines so that two longitudinal rows of unequal rectangles (one large, one very
small) are formed.

ectocommensal: living on the surface ofand in close association with another organism of a
different species where neither has an obvious effect on the other, cf. endocommensal.

elevation: of diatoms, Family Eupodiscaceae, a raised portion of a valve wall, not projecting
laterally beyond the valve margin, which may bear some special structure; otherwise
with the same structure as valve.

"Eltanin": U.S.A. Oceanographic Research Ship Eltanin, Cruise 38, Southern Ocean, south
of Australia, 1969.

EM: Elecron Microscope/Microscopy; see SEM and TEM.

encystment: process of forming a protective stage in a life history in which an organism
surrounds itself with a cyst wall.

endocommensal: living inside another organism of a different species where neither has an
obvious effect on the other, cf. ectocommensal.

475
 GLOSSARY

endoral membrane: of ciliates, a row of cilia extending on the bottom and right border of
the oral cavity.

endostyle: of loricate peritrich ciliates, an internal stalk connecting a zooid to the bottom of
lorica or (when present) to the septum (an extra layer of material forming internal lining
in lorica).

endosymbiont: an organism living within a dissimilar organism in a relationship in which

both partners benefit, adj. endosymbiotic.

endotheca: the inner layer of coccoliths of a dithecate coccolithophore. adj. endothecal.

entoecism (= inquilism): of ciliates, an intimate (transitional) association between two

organisms (usually animals) in which one partner lives within the host body to obtain
shelter but without parasitising it. adj. entoecic.

epibiotic: an organism that grows attached to the outside of another living organism.

epiphytic: a plant that grows attached to the outside of another plant, using it only for
support, but doing it no harm.

episome: see epitheca.

epitheca (= episome): ofdinoflagellates, the anterior part a of a dinoflagellate cell above the
cingulum. cf. hypotheca.

epizoic: an organism that grows attached to the outer surface of an animal.

equant: equal in size and form.

ER: Endoplasmic Reticulum, a series of interconnected, flattened, subcellular cavities.

"Erebus": Royal Society (London) Expedition Ship Erebus, Antarctic expeditions of 1839—43,
commanded by James Clarke Ross.
eukaryote: any cell or organism comprised of cells that possess a membrane-bound nucleus,
several chromosomes, cellular organelles and accomplishes cell division by mitosis and
meiosis. adj. eukaryotic; cf. prokaryote.

euglenoid: a member of the Class Euglenophyceae.

euhedral: of crystals, characterised by well-developed crystal faces (external surfaces).

exotheca: the outer layer of coccoliths of a dithecate coccolithophore. adj. exothecal.

extrusome: of ciliates, a membrane-bound, subpellicularly located extrusible body of

various, probably non-homologous types (e.g. cyrtocysts, mucocysts, pexicysts,
toxicysts, trichocysts).

extrusome warts: of ciliates, Genus Loxophyllum, protruding, wart-like structures along

dorsal margin containing groups of extrusomes (toxicysts).

eyespot (= stigma): a pigmented, light-sensitive organelle.

fascia: of diatoms, Family Achnanthaceae, an extension of the central area forming a hyaline
(unperforated) band extending the valve, i.e. transapically.

fasciculate areolation: of diatoms, valve striae (and, therefore, areolae) grouped in discrete
sectors (fascicles).

fenestra: a small, window-like perforation, pi. fenestrae; adj. fenestrated.

Feulgen-stained: material treated with a histological stain selective for DNA.

FIBEX: First International Biological Experiment; international research cruise, 1981,
Southern Ocean between Mawson and Davis Stations.

476
 GLOSSARY

fibula (= keel punctum): of diatoms. Family Bacillariaceae, an internal bridge of silica

subtending the raphe, pi. fibulae.

filopodium: of amoebae, a filiform pseudopodium with a pointed tip, capable of rather rapid
movement, used for locomotion and food uptake.
filose: of amoebae, of the Class Filosea.

filter feeding: atype of feeding in which suspended particles are concentrated and ingested
by means of cilia or flagella directing a current of water to the cell, and a filter to trap
and concentrate the particles before enclosure within a food vacuole.

flagellum: a fine, whip-like process associated with locomotion, normally with the typical
9><2 + 2 microtubular arrangement of the axonemes. pi. flagella; adj. flagellar,
flagellate, flagellated.

flagellar pore: of dinoflagellates, a pore through which a flagellum emerges.

flimmer hairs: of chrysophytes, minute tubular hairs along all or part of the flagellar axis.

food vacuole: a fluid-filled space within the cell cytoplasm delimited by a membrane, the
repository for food particles ingested by phagocytosis.

foot pole: see antapical pole; cf. apical pole, head pole.

footprint scale: of prasinophytes, a minute, organic body scale in the shape of a human
footprint.

foramen (= aperture): of diatoms, an opening or passage through the constriction at the

valve surface opposite the velum, pi. foramina.

Formvar: a material for producing very thin films for supporting material on electron
microscope grids.

frontal cirri: of spirotrich ciliates, near the anterior end of a cell, often slightly enlarged.

frontal plate: of ciliates, a non-ciliferous, often transversely truncate apical area.

frontoterminal cirri: of urostylid ciliates, a short row of cirri that originates during
morphogenesis near the transverse cirri and migrates anteriorly close to the right end of
the adoral zone of membranelles.

frontoventral cirri: of spirotrich ciliates, a group of cirri between the anterior portion of the
right-marginal cirral row (or the right body margin in euplotids) and the paroral membrane.

fusiform: swollen at the middle and tapering towards each end; narrowly ellipsoidal.

frustule: the silica-containing wall of a diatom.

gamete: a haploid cell that fuses with another to produce a diploid zygote, adj. gametic.

"Gauss": German South Polar Expedition aboard the Gauss, 1901-03, Davis Sea (see Heiden
&Kolbe, 1928).

genuflexed: bent like a knee.

girdle: of diatoms, part of a frustule between the upper and lower valves; of strombidiid
ciliates, referring to transverse orientation.

girdle band: see intercalary band, copula.

girdle view: of diatoms, a view of the cell with the valves seen in profile and the girdle seen
in face view.

gullet: of cryptophytes, the buccal cavity at the cell apex from where the two flagella arise.

gyre: a coil.

477
 GLOSSARY

haploid: a vegetative or gametic organism in which the nucleus contains one set of
chromosomes, abbrev. nlln.

haptonema: of haptophytes, an organelle formed as a microtubular appendage, serving as a

holdfast or for crawling or food uptake.

haptophyte: a member of the Class Haptophyceae.

head pole: see apical pole; cf. antapical pole, foot pole.

helicoid: spiral.

helicolith: of coccolithophores, a placolith with a spiral pattern and marginal flange which
overlaps adjacent helicoliths (e.g. Helicosphaera carteri).

helictoglossa: of diatoms, Family Bacillariaceae, an inwardly projecting structure

resembling a rolled tongue, located at the distal end of a raphe on the internal valve
surface.

herbivorous: of an organism that feeds exclusively on plant material, n. herbivory.

heterococcolith: of coccolithophores, a calcite scale (coccolith) built up of complex, calcite

crystal units of various shapes and sizes, cf. holococcolith.

heterodynamic flagella: flagella that have independent patterns of beating, cf. homodynamic
flagella.

heterokont: ofdinoflagellates, biflagellate, with one smooth flagellum and the other covered
with mastigonemes.

heteropolar: of diatoms, cells having morphologically different poles (apices), cf. isopolar.

heterotroph (-y): an organism that is dependent on obtaining organic food from the
environment due to its inability to synthesise organic material, adj. heterotrophic; cf.
autotroph, mixotroph.

heterovalvate: of diatoms, cells having morphologically different valves, cf. isovalvate.

holdfast: the basal part of an organism attaching it to the substratum.

holococcolith: of coccolithophores, a calcite scale (coccolith) built up of discrete calcite

crystallites (often rhomboidal), each enclosed by a thin organic envelope, cf.

heterococcolith.

holotrichous: of ciliates, having the entire body completely and ±uniformly ciliated.

homodynamic flagella: flagella that have the same pattern of beating, cf. heterodynamic
flagella.

horn: of diatoms, Family Asterolampraceae, a long, narrow elevation; of dinoflagellates,

prominent extensions of the cytoplasm covered by thecal plates of epi- or hypotheca.

hyaline: colourless and transparent.

hyaline band: of diatoms, an element of a girdle lacking perforations.

hyaline ray: of diatoms, Family Asterolampraceae, raised openings in the valve face, open
to the interior of the valve by elongate, slit-like openings, open to the exterior through
holes at the marginal ends of the rays.

hyposome: see hypotheca.

hypotheca (= hyposome): ofdinoflagellates, posterior part of a cell bearing one longitudinal

and one transverse flagellum; of diatoms, the lower part of a frustule. cf. epitheca.

infraciliature: of ciliates, the total assembly of somatic and oral basal bodies and their
associated subpellicular microfibrillar and microtubular structures.

478
 GLOSSARY

inner seta: of diatoms, an intercalary seta or a seta occurring within the chain of cells, pi.

setae.

intercalary band (= copula, girdle band): of diatoms, a single element of the girdle; the
element nearest to the valve, different in structure from elements farther away.

intercalary plate: of dinoflagellates, thecal plates not in contact with the apex or the
cingulum (for anterior intercalary plates), or with the antapex or the cingulum (for
posterior intercalary plates).

interstria: of diatoms, the non-perforate, siliceous strips between two striae, pi. interstriae;

adj. interstrial.

invagination: inpushing of a layer of cells; or a concave structure (e.g. cell surface, oral

area) in unicellular organisms.

in vivo: in reference to a living organism.

interstrial costa, interstrial membrane: of diatoms, a siliceous rib on a valve surface in the

space between two striae.

investment: a covering or coating.

isopolar: of diatoms, cells having morphologically identical poles (apices), cf. heteropolar.

isovalvate: of diatoms, cells having morphologically identical valves, cf. heterovalvate.

keel: see raphe keel.

keel punctum: see fibula.

KERFIX: a permanent sampling station during 1992-95, south-west of Kerguelen Is., near
the Polar Front at 50°40'S 68°25'E.

and flagellates, basal apparatus of cilia and flagella, consisting of at least

kinetid: of ciliates
one kinetosome with all associated rootlets (monokinetid), but also of two (dikinetid) or
more kinetosomes (polykinetid).
kinetofragmon: of ciliates, Family Kryoprorodontidae, a short (curved) row of somatic cilia

transversely on lateral side of body, very likely a reduced spiral kinety.

kinetosome (= basal body): of ciliates, a subpellicularly located basal termination of cilia,

cylindrical and composed of 9 peripheral microtubular triplets; may be non-ciliferous.

kinety: of ciliates, a structurally and functionally integrated row of single or paired somatic
kinetids, often running longitudinally.

kummerform: in reference to the final chamber of a whorl being reduced in size.

labiate process (= rimoportula): of diatoms, a tube or an opening through the valve wall
with an internal flattened tube or a longitudinal slit surrounded by two lips.

laevogyre: of coccolithophores elements and sutures that curve to the

, left when traced
radially outward, e.g. in Calcidiscus leptoporus.

lamina: of ciliates, a layer, adj. luminal

lanceolate: narrow with curved sides tapering to a pointed end.

lateral: of, at, towards or from the side.

lath/lath-like element: of coccolithophores, a thin strip of crystalline structure.

left: left-hand (view or side) in relation to dorsal surface.

479
 GLOSSARY

life cycle: the continuum of phases and cell types in the growth and reproduction of an
organism.

ligula: a small, tongue-like projection on diatom copulae. pi. ligulae.

limuliis scale: of prasinophytes, a minute, organic body scale in the shape of a horseshoe
crab.

linking spines: of diatoms, Family Fragilariaceae, marginal spines that link sibling valves
in a chain.

list (= wing, flange): a membranous extension of an armoured dinoflagellate.

LM: light microscope/microscopy.

lobopodium: in amoebae, a lobe-like pseudopodium with a rounded termination, used for

locomotion and food uptake.

loculus/loculate areola: of diatoms, an areola (pore) markedly constricted at one surface of

the valve and occluded by a velum at the other.

lorica: of choanoflagellates, siliceous investment surrounding a cell body; of ciliates, not only
of siliceous material but also, e.g. proteinaceous or of mucopolysaccharides, adj. loricate.

lorica chamber: a cavity created by a choanoflagellate lorica.

macronucleus: of ciliates and foraminiferans, a large, somatic nucleus that determines the
phenotype of the cell; found in cells with nuclear dimorphism, cf. micronucleus.

macrozoospore: of the haptophyte Phaeocystis, a comparatively large, motile, flagellated

spore capable of vegetative multiplication.

mammiform: swollen, with a small, nipple-like projection.

marginal cirri: of spirotrich ciliates, one to few rows of cirri along lateral cell margins.

marginal ridge: of diatoms, a ridge between the valve face and the valve mantle, continuous
or interrupted, perforated or solid.

mastigoneme (= flimmer): a hair-like projection of the flagellar shaft.

median cingulum (= equatorial girdle): of dinoflagellates, a cingulum located approx-

imately at the midpoint of the cell.

membrane: a thin sheet of organic material in protists, plants and animals.

membranelle (= polykinety): of ciliates, a compound structure comprising many cilia, and

associated with the mouth of a ciliate; present either in groups of 3 or as a band of many
more.

mesoplankton: plankton organisms in the size range 200-2000 urn. adj. mesoplanktonic.

mesostyle: of loricate peritrich ciliates, a short, broad and non-contractile internal middle
stalk between septum (extra layer in lorica) and base of lorica.

metaboly: mobility and the ability to change the shape of the cell.

"Meteor": Indian Ocean Expedition of R.V. Meteor, 1964-65 (see Simonsen, 1974).

micron: see micrometre.

6
micrometre = micron):
( a unit of length equivalent to 10" of a metre, abbrev. um.

micronucleus: of ciliates and foraminiferans, a small, germline nucleus that shows little or
no gene expression, but with the potential to undergo meiosis; found in cells exhibiting

nuclear dimorphism, cf. macronucleus.

microplankton: plankton organisms in the size range 20-200 um. adj. microplanktonic.

480
 GLOSSARY

microtubule: a thin, proteinaceous tubule acting as a supporting, cytoskeletal element within

protist cells.

microzoospore: of the haptophyte Phaeocystis, a comparatively small, motile, flagellated

spore capable of vegetative multiplication.

midventral cirri: of spirotrich ciliates, a usually longitudinal row of cirral pairs in a zig-zag
arrangement ±along the median of a cell.

mitochondrion: a cylindrical, subcellular organelle found in eukaryotes, containing some

DNA and the enzymes required for metabolic processes, pi. mitochondria.

mixotroph (-y): an autotrophic organism taking up organic substances for supplementary

nutrition (e.g. by phagotrophy), or a heterotroph with ingested chloroplasts supplem-
enting nutrition, adj. mixotrophic.

monad: the flagellated stage in the life cycle of protozoa and algae.

monokinetid: of ciliates, the derived formation of unpaired individual kinetids.

monostromatic: single-layered, usually only one cell thick.

morphotype: an organism with a variation in form or structure from the type specimen, but
considered to be circumscribed within the same species.

motile: moving, e.g. by swimming, gliding, crawling, kicking or pulsating.

mucocyst: of ciliates, a type of extrusome, mucilaginous, with a protective function against

(for example) toxins, and possibly involved in cyst formation.

murolith: a heterococcolith with an elevated rim but without a well-developed shield,

e.g. Syracosphaera pulchra.

naked: uncovered, e.g. a choanoflagellate or ciliate without a lorica, a dinoflagellate without

thecal plates, or an amoeba without a test.
9
nanometre: a unit of length equivalent to 10~ of a metre, abbrev. nm.
nanoplankton: plankton organisms in the size range 2-20 urn. adj. nanoplanktonic.

narrow girdle view: a frustule seen from the narrow side.

nematodesma (= cytopharyngeal rod): of ciliates, a rod-like, microtubular bundle arising

from the distal side of neighbouring kinetosomes and cross-linked, often in a hexagonal
pattern, pi. nematodesmata.

nematocyst: of dinoflagellates, an ejectile organelle, often arranged radially or subradially

in the cell (e.g. in Polykrikos).

nilas ice: a very thin, grey, elastic type of ice with a matt surface which is readily bent by
wave action.

non-ciliferous: without cilia.

non-contractile: not able to contract.

non-motile: not moving.

nucleolus: an organelle, containing rRNA, within a eukaryotic nucleus.

nucleus: a membrane-bound organelle in eukaryotic cells containing the chromosomes

whose genes control the structure of proteins within the cell.
nuclear dualism: of ciliates, the simultaneous presence of somatic (macro-) and generative
(micro-) nuclei within a cell.

481
 GLOSSARY

ocellus: of diatoms, a small, siliceous thickening in the valve wall, usually with a thickened,
structureless surrounding an area of small pores (e.g. in Eucampia); of
rim
dinoflagellates, modified region of the chloroplast resembling a light-perceiving
a
organelle, typically located in the left side of the cell.

occluded process: of diatoms, Family Thalassiosiraceae, a hollow, external tube not

penetrating the valve wall, sometimes at the top of an areola.

operculate strutted process: of diatoms, Family Thalassiosiraceae, a strutted process

penetrating a diatom valve, but which also has a small covering, or operculum,
occluding the satellite pores around the base of each process.

oral apparatus (= buccal apparatus, mouth): of flagellates and ciliates, infraciliature

located in or associated with the oral area.

oral cone: of ciliates, an everted, cytopharyngeal apparatus at anterior the end of a cell body.

otarium (= wing): of diatoms, Family Rhizosoleniaceae, one of a pair of membranous costae

that occur opposite each other at or near the base of the external process, pi. otaria.

oxic: containing oxygen, cf. anoxic.

palmella stage: of haptophytes, Order Prymnesiales, a stage during which flagella are
discarded, and cells proliferate within a mucilaginous matrix, adj. palmelloid.

panduriform: fiddle-shaped.

papilla: a small projection from the surface, adj. papillate.

pappolith: of coccolithophores, a heterococcolith with a rim in the form of a narrow,

slightly flanged wall composed of elements of two alternating type, one relatively small
and confined to the proximal part of the wall, and the other vertically expanded, e.g. in
Papposphaera spp.

paracingulum: a cingular area in dinoflagellate cysts.

parietal: borne or located on the wall.

paroral: of ciliates, at or along the right side of the buccal cavity. Also used as a noun for
paroral membrane, i.e. a ciliary organelle lying along the right side of the buccal cavity.

pedicel: a type of holdfast organelle.

peduncle: a cytoplasmic appendage located near flagellar pores in some photosynthetic and
non-photosynthetic dinoflagellates, associated with phagotrophy.

pellicle: a rigid or semi-rigid investment of some dinoflagellates and all ciliates. adj.
pellicular.

peniculus: a type of ciliary membranelle positioned along the left wall in the buccal cavity
of some oligohymenophorean ciliates. pi. peniculi.

pennate: referring to diatoms of the Order Pennales.

perioral: around the oral area.

peristomial field: of ciliates, generally refers to the entire, expansive, "everted" oral area
with its encircling adoral zone of membranelles, etc.

peritrich: Subclass Peritrichia of Class Oligohymenophorea, a type of ciliate which, in the

adult stage, has a characteristically inverted bell- or goblet-shaped or conical-cylindrical
body, a conspicuous buccal ciliature at the apical pole, the somatic ciliature reduced to a
subequatorial band and often a stalk (contractile or acontractile) or adhesive disc at the
antapical pole; many species are sedentary (Order Sessilida) and form colonies.

pervalvar axis: the axis through the centre point of two valves.

482
 GLOSSARY

pexicyst: of ciliates. Family Didiniidae, a type of toxicyst-like extrusome which, on

discharge, adheres to the surface of prey without subsequent penetration.

petaloid: resembling the petal of a flower.

phagocytosis: ingestion of particulate food.

phagotroph (-y): any heterotrophic organism that ingests or absorbs nutrients as solid
particles from itssurroundings, adj. phagotrophic.

pharyngeal basket: of ciliates, a tubular cytopharyngeal apparatus, often curved, with walls
strengthened by longitudinally arranged nematodesmata. Depending on the origin of the
microtubules, a cyrtos- or rhabdos-type can be distinguished.

pharyngeal plug (= oral bulge): of ciliates, a ±bulge-like structure on the apical cell pole in

the centre of which lies the cytostome.

phytoflagellate: a photosynthetic, flagellated organism.

phytoplankton: that part of the plankton made up of autotrophic organisms.

picoplankton: plankton organisms in the size range 0.2-2 urn. adj. picoplanktonic.

pigment: coloured material in organisms, often associated with photosynthesis.

pinocytosis: of dinoflagellates, the active engulfing of small particles or liquids from prey
cells.

placolith: of coccolithophores, a heterococcolith composed of an upper and lower shield of

radial elements, interconnected by a central column, e.g. in Calcidiscus leptoporus.

plankton: organisms drifting with the current, generally inhabiting the water column of an
ocean, sea or lake; many planktonic organisms are capable of locomotion but are not
sufficiently strong swimmers to move against the current, adj. planktonic.

plate formula: see tabulation.

pleura (= connecting band): of diatoms, an element in the middle of the girdle when
intercalary bands are present; any element in the absence of intercalary bands, pi. pleurae.

podite: see stylus.

pomiform: apple-shaped.
pore/poroid: of diatoms and dinoflagellates, an opening or channel in a theca or cell wall.

poroid areola: of diatoms, an areola not markedly constricted at one surface of the valve.

"Porquoi Pas?": Deuxieme Expedition Antarctique Francaise, 1908-10, commanded by Dr

Jean Charcot, on board the Porquoi Pas?.

posterior: that part of the organism which proceeds last in forward movement.

postmedian cingulum: of dinoflagellates, a cingulum located below the midpoint of the cell.

ppm: parts per million.

precyst: of testate amoebae, a short-term, dormant stage in the life cycle, enclosed by a
relatively thin membrane (not a cyst wall), and with an internal diaphragm closing the
aperture of the test.

prasinophyte: a member of the Class Prasinophyceae.

premedian cingulum: of dinoflagellates, a cingulum located above the midpoint of the cell.

preoral: of ciliates, anterior to the cytostome.

pretransverse cirri: of spirotrich ciliates, usually few ventral cirri immediately anterior to
the larger transverse cirri.

483
 GLOSSARY

proboscis: of diatoms, Family Rhizosoleniaceae, a permanent, elongate part of the valve

with a truncate tip, considerably thicker than a flagellum (e.g. in Proboscia); the distal
part fits into a groove on the adjacent valve of linked cells; of ciliates, Family
Didiniidae, see oral cone.

prokaryote: any cell lacking a true nucleus, usually with only one chromosome, lacking
cellular organelles and accomplishing cell division by binary fission, adj. prokaryotic;
cf. eukaryote.

process: of diatoms, a homogeneously silicified projection of a cell wall; in Family

Rhizosoleniaceae, the process is spine-like.

prolate: lengthened, drawn out towards the poles.

protargol: protein silver impregnation of ciliates to stain the infraciliature (mainly

kinetosomes, occasionally fibres), often nuclei and sometimes extrusomes, used to
identify the ciliary structures.

protist: a member of the Kingdom Protista.

protoplasm: the living contents of a cell, i.e. the cytoplasm and nucleus, adj. protoplasmic.

protoplast: the main constituents of a cell; a cell excluding the cell wall.

protozoa: members of the Phylum Protozoa, a term widely used to refer to heterotrophic
flagellates, ciliates and amoebae.

pseudocellus: of diatoms, Family Eupodiscaceae, an area of small, unoccluded pores at the

cell apices which grade into the areolae on the valve face. cf. ocellus.

pseudonodulus: of diatoms, a solitary, marginal or submarginal structure evident as an open

hole or an area covered by densely packed smaller areolae by LM. The principle
diagnostic feature of Family Hemidiscaceae.

pseudopod/pseudopodium/pseudopodial projections: of Class Heliozea, and also of

amoebae, temporary extensions of the cell surface without defined shape, functioning in
locomotion and/or food uptake.

pseudoraphe: see raphe sternum.

pseudostome: of testate amoebae, an aperture in the test through which the pseudopodia
protrude.

ptychocyliform: of tintinnid ciliates, lorica shape as in genus Ptychocylis.

punctate: dotted, marked with dots. n. puncta; pi. punctae.

pyrenoid: a region of the chloroplast involved in starch formation in certain protists.

pyriform: pear-shaped.

raphe: of diatoms, Family Bacillariaceae, one or two longitudinal slits through the valve
wall of raphid pennate diatoms possibly involved in the secretion of mucus for
locomotion, cf. raphe sternum.

raphe canal: of diatoms, Family Bacillariaceae, a space on the inner side of the raphe cut
off from the rest of the interior of the frustule.

raphe fins: of diatoms, Family Naviculaceae, small, paired, external, vane-like (or blade- or
plate-like) siliceous structures arising from the raphe sternum.

raphe fissure: of diatoms, a narrowing opening of the raphe slit; terminal fissures at the

valve ends can be hooked, Y- or T-shaped.

raphe keel: of diatoms, Family Naviculaceae, an elevation of the valve which bears and
raises the raphe; also possessing specialised supporting structures (fibulae).

#
484
 GLOSSARY

raphe slit: of diatoms, a longitudinal slit in the valve of some pennate diatoms, typically
along or near the midline of the valve.

raphe sternum (= pseudoraphe): of diatoms, an elongate part of the valve where areolae are
sparse or absent and which is often thickened towards the perivalvar axis. cf. raphe.

raphe valve: of diatoms. Order Pennales, the valve bearing the raphe.

reniform: kidney-shaped.

reticulum: a network, adj. reticulate.

rhabdos: of ciliates, a type of tubular cytopharyngeal apparatus; not curved, may be eversible,
with walls strengthened by longitudinally arranged nematodesmata. Derived from
circumoral kinetosomes and lined with extensions of transverse microtubules; often
contains toxicysts.

rhombic (-oid): having the shape of an oblique, equilateral parallelogram.

ribs: of dinoflagellates, Family Protoperidiniaceae, supports for sulcal or cingular lists.

ribbons: of diatoms, cells joined by abutting valve faces (Fragilariopsis) or by marginal

linking spines (Synedropsis).

right: right-hand (view or side) in relation to the dorsal surface.

rimoportula: see labiate process.

rostrate: beak-like.

rota: of diatoms, a solid disc attached by spokes to the surrounding silica framework of a
velum, pi. rotae.

rRNA: ribosomal RNA (ribonucleic acid), a nucleic acid within the cells of living organisms
that is a vital component of protein synthesis.

saprobe: an organism that feeds on soluble organic nutrients of dead or dying plants or
animals, adj. saprobic.

satellite pores: of diatoms, Family Thalassiosiraceae, small pores associated with strutted
processes, penetrating a diatom valve; pores are sometimes occluded by an operculum
(operculate strutted process).

scale: a calcified or silicified, plate-like outgrowth.

scutica: of ciliates, Subclass Scuticociliatia of Class Oligohymenophorea, usually a small,

hook-like or whiplash group of often non-ciliferous kinetosomes posterior to the oral
apparatus; during cell division, part of the new oral structures develops from this.

segment (= band): of diatoms, a single element of the girdle.

SEM: scanning electron microscope/microscopy.

septum: of diatoms, a sheet or ridge in the valvar plane projecting from a girdle band into
the interior of the frustule, often with several openings.

seta: a hollow, bristle-like structure, often chitinous (in e.g. Chaetoceros). pi. setae.

sessile: fixed to a substratum, e.g. by a stalk or lorica.

sexual reproduction: involving the production of haploid gametes that fuse to produce a
diploid zygote.

sheath: a loose covering, usually of mucus (as in the lorica sheath of Acanthocorbis
unguiculata).

485
 GLOSSARY

SIBEX: Second International BIOMASS Experiment 1985, international research cruise on

board R.V. Nella Dan, Indian Ocean sector of the Antarctic.

sigmoid: curved like the letter S.

siliceous: of, relating to, or incorporating silica.

silicoflagellate: a member of the Class Dictyophyceae.

silverline system: ofciliates, the entire system of pellicular or cortical structures, organelles
or markings that can be revealed by silver nitrate impregnation; often, however, referring
only to the striated or reticulate system of 'lines' (fibres); structures are possibly
involved in morphogenesis or conduction of stimuli.

skeleton: any structure in an organism that maintains its shape and supports the structures
associated with the body, e.g. silicoflagellates have a siliceous skeleton.

somatic: relating to the body surface, or to non-reproductive structures or activities; in

ciliates, relating to all parts of the body not associated with the oral structures.

spathulate: spoon-shaped; broad at the tip and narrowed towards the base.

spicule: a delicate, pointed structure external to the cell, invariably excreted by the cell;
more delicate than a spine.

spine: a solid protuberance or sharp, pointed structure, either part of a cell or secreted by it.

spine scale: a plate-like, secreted structure with a spine-like protrusion, usually siliceous.

spinule: a small spine.

spore: usually, a drought-resistant propagule and infection-inducing stage in the life cycle of
several parasitic protozoa.

stellate: star-shaped.

stepped chains: of diatoms, chains with overlapping cell ends (as in Pseudonitzschia).

sternum: see raphe sternum.

stigma: see eyespot.

stigma: of diatoms, an isolated pore, typically in the central area of a valve face.

stria: of diatoms, surface ornamentation of areolae, pores of an alveolus appearing as lines

or ridges. Orientation of striae is useful in identification, e.g. parallel, radiating, linear or
convergent, pi. striae; adj. striate.

strutted process: of diatoms, Family Thalassiosiraceae, a tubular process penetrating a

diatom valve, through which a thread of organic material (mostly chitin) is extruded, see
trifultate. operculate strutted process; cf. occluded process.

styliform: needle-like.

stylus (= podite): of ciliates, Class Phyllopharyngea, a conical, foot-like appendage

projecting from the posterior ventral surface, through which a mucus-like filament can be
secreted for temporary attachment to a substratum.

subpellicular platelet layer: of strombidiid ciliates, a rigid layer of small, polygonal

polysaccharide plaques (skeletal elements) subpellicularly in the posterior portion (often
half) of a cell.

suctoria: ofciliates, Subclass Suctoria of Class Phyllopharyngea, a type of ciliate which, in

the adult stage, is generally sedentary, lacks cilia, feeds by means of suctorial tentacles,
and often has a non-contractile stalk; in the swarmer stage, is ciliferous and mobile.

sulcus: a longitudinal area on the ventral surface of a dinoflagellate cell that forms the
furrow housing the longitudinal flagellum.

486
 GLOSSARY

suture: of dinoflagellates, a visible, linear boundary between thecal plates.

swarmer: a freely motile dispersal stage in the life cycle of a number of sessile ciliates (e.g.
suctorians, peritrichs).

symbiont: an organism living in a state of symbiosis with another, a relationship in which

both partners benefit.

synhymenium: of nassophorid ciliates, a hypostomial flange, perioral ciliature; a band of

('membrane') slightly posterior to oral area, ±traversing helically the ventral
ciliature
side and sometimes extending to the dorsal side, composed of closely set kinetal
segments (mono-, di- or polykinetids).

systematics: the area of biology dealing with the classification, naming (nomenclature) and
evolution of organisms.

tabulation (= plate formula): the formula describing thecal plate orientation in armoured
dinoflagellates.

TEM: transmission electron microscope/microscopy.

tentacle: of suctorians, a long, tubular cell projection, usually stiffened by microtubular

bundles and equipped with haptocysts at the tip, extensible and retractable, used in prey
capture and food uptake.

terminal nodule: of diatoms, Family Bacillariaceae, a thickening at the apical end of the
raphe.

terminal seta: of diatoms, a seta on the end valves of a chain.

"Terror": Royal Society (London) Expedition Ship Terror, Antarctic expeditions of 1 839—
43, commanded by Francis Crozier.

test: a general term for a rigid shell or lorica around an organism, adj. testate.

theca: an extracellular covering, e.g. test, valve or frustule, enclosing a cell. pi. thecae.

thecal plates: plates of armoured dinoflagellates composed of cellulose or other

polysaccharide microfibrils.

thecostyle: of suctorian ciliates, a posterior, stalk-like extension of the lorica.

thigmotactic: of ciliates, adhering; also referring to specialised, somatic cilia that are
modified to serve a tactile or adhering function.

thylakoid: a thin-layered plate within a chloroplast.

toxicyst: of ciliates, a type of toxic extrusome, used in prey capture.

transapical axis: the third axis of a bilateral diatom, cf. apical axis, pervalvar axis.

transapical plane: perpendicular to the transapical axis.

transverse cirri: of spirotrich ciliates, a group of often enlarged cirri in the posterior quarter
of a cell, frequently in an oblique, J-shaped row.

trichocyst: of dinoflagellates and ciliates, an ejectile organelle.

trichocyst pore: of dinoflagellates, an opening or channel in the theca, possibly involved in

pinocytosis, extrusion of trichocysts and mucocysts and other active processes.

trifultate strutted process: of diatoms, Family Thalassiosiraceae, a strutted process,

penetrating a diatom valve, with a long internal tube with three columnar supports.

triradiate: radiating in three directions.

triundulate: with three undulations or bulges.

487
 GLOSSARY

trochospiral: of foraminiferans relating to chambers that are spirally coiled.

,

trophont: of dilates, an adult trophic and growing stage of the life cycle.

truncate: ending abruptly, as if cut off.

tubular process: of diatoms, a simple tube penetrating the valve wall.

tychopelagic: of diatoms, relating to the bottom of a substratum, e.g. ice, until detached into
the surrounding water.

umbilicus: offoraminiferans having a small, central depression or hollow,

, adj. umbilicate.

unarmoured (= naked, athecate): of dinoflagellates, cells without an identifiable theca or

cell wall; while they do not have apical pore complexes, there may be an apical groove.
Some species with thin thecal plates and apical grooves are considered to be transitional
between unarmoured and armoured cells.

vacuole: a membrane-enclosed compartment in the cell, e.g. food vacuole, contractile vacuole.

valvar plane: a plane of division parallel to the valve surface.

valve: two opposing halves of theca or test.

valve face: the entire diatom valve surface, surrounded by the mantle.

valve mantle: marginal parts of a diatom valve, set off from the valve face at an angle.

valve view: a diatom frustule seen from top or bottom.

valvocopula: of diatoms, an intercalary girdle band adjacent to a valve.

varimorphic: coccoliths on a coccosphere showing a continuous range of morphology

between two end member morphologies, e.g. in Papposphaera borealis.

velum: of diatoms, a thin, perforated layer of silica across an areola; of dinoflagellates, a

flat, cellular projection, especially in heterotrophic dinoflagellates, involved in
locomotion and food uptake.

ventral: the underside of an organism; in dinoflagellates, the ventral side is identified by the
presence of the sulcus; in dilates, by the position of the cytostome, except when this is
apical, then opposite to brosse (exception: suctorians); in testate amoebae, by the
position of the pseudostome.

ventral ridge (= ventral flange): of dinoflagellates, an identifiable ridge on the right side of
the sulcal intrusion onto the epitheca.

vermiculate: with raised, worm-like markings, e.g. on the thecal plates of armoured
dinoflagellates.

zig-zag or stellate colonies: of diatoms, cells joined in star-shaped colonies by mucilage

pads (e.g. in Thalassionema).

zooid: of dinoflagellates and ciliates, a motile cell or body produced by an organism or

individual member in a colony.

488
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With permission from the Journal of Phycology:

Marchant, H.J., Buck, K.R., Garrison, D.[L.] & Thomsen, H.A. (1989), Mantoniella in Antarctic waters
including the description of M. antarctica sp. nov. (Prasinophyceae), J. Phycol. 25: 167-174. Fig. 1

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With permission from Kluwer Academic Publishers:

Everitt, D.A. & Thomas, D.P. (1986), Observations of seasonal changes in diatoms at inshore localities
near Davis Station, East Antarctica, Hydrobiologia 139: 3-12. Fig. 2F [Fig. 2.67c], Fig. 3B [Fig. 2.6c],
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Perrin, R., Lu, P. & Marchant, H.J. (1987), Seasonal variation in marine phytoplankton and ice algae at

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With permission from the Marine Biological Association of the United Kingdom and Cambridge
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strobilidiine ciliates (Ciliophora: Choreotrichida) from the Isles of Shoals, Gulf of Maine, J. Mar. Biol.
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Lynn, D.H., Montagnes, D.J.S. & Small, E.B. (1988), Taxonomic descriptions of some conspicuous
species in the family Strombidiidae (Ciliophora: Oligotrichida) from the Isles of Shoals, Gulf of Maine,
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With permission from Museo Argentino de Ciencias Naturales Bernardino Rivadavia:

Balech, E. (1971), Microplancton de la campafia oceanografica Productividad III, Revta Mus. Argent.
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With permission from Museo de La Plata:

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With permission from the National Institute of Polar Research:

Hada, Y. (1970), The protozoan plankton of the Antarctic and Subantarctic Seas, JARE Sci. Rep., Ser.
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Takahashi, E. (1981), Loricate and scale bearing protists from Liitzow-Holm Bay Antarctica. I. Species
of the Acanthoicidae and the Centrohelia found at a site selected in the fast ice, Antarc. Rec. 73: 1-22.
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Serrano, S., Sola, A., Guinea, A., Arregui, L. & Fernandez-Galiano, D. (1992), Cytoskeleton of Euplotes
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With permission from the Nencki Institute of Experimental Biology, Polish Academy of Sciences:

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Agatha, S., Wilbert, N., Spindler, M. & Elbrachter, M. (1990), Euplotide ciliates in sea ice of the
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Fig. 17a, c, d [Fig. 14.21a-c], Fig. 15a, c, d, f [Fig. 14.26a-d], Fig. 12a, h, j [Fig. 14.85a-c], Fig. 3a, f,

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Agatha, S., Spindler, M. & Wilbert, N. (1993), Ciliated protozoa (Ciliophora) from Arctic sea ice, Acta
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Song, W. & Wilbert, N. (2002), Faunistic studies on marine ciliates from the Antarctic benthic area,
including descriptions of one epizoic form, 6 new species and, 2 new genera (Protozoa: Ciliophora),
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With permission from the Royal Society of New Zealand (New Zealand Journal of Botany):

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With permission from OO Landesmuseum, Biologiezentrum:

Petz, W., Song, W. & Wilbert, N. (1995), Taxonomy and ecology of the ciliate fauna (Protozoa,
Ciliophora) in the endopagial and pelagial of the Weddell Sea, Antarctica, Stapfla 40: 1-223. Fig. 31a, b
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b, g [Fig. 14.23a-d], Fig. 55a, b, d [Fig. 14.24a-c], Fig. 56a-c [Fig. 14.25a-c], Fig. 57a, d, e, f [Fig.
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14.29a-d], Figs 43a-e, 63 [Fig. 14.41a-e], Fig. 41a, d [Fig. 14.78a, b], Fig. 44 [Fig. 14.45], Fig. 45a-d
[Fig. 14.58a-d], Fig. 46a-d, f [Fig. 14.59a-d, f], Fig. 42a-c [Fig. 14.80a-c], Fig. 40a, c, d [Fig. 14.81a-
c], Fig. 50a-c [Fig. 14.83a-c], Fig. 47a-c [Fig. 14.86a-c], Fig. 49a-c [Fig. 14.87a-c], Fig. 48a-c [Fig
14.88a-c], Fig. 39a, b, d, e [Fig. 14.92a-d], Fig. 33a-c [Fig. 14.94a-c], Fig. 34a, c, d [Fig. 14.96a-c]
Fig. 35a-c [Fig. 14.101a-c], Fig. 36a, c, d [Fig. 14.102a-c], Fig. 37a, b [Fig. 14.106b, c], Fig. 38a-c
[Fig. 14.107a-c], Fig. 6a, b [Fig. 14.1 10a, b], Fig. 4a, c, d [Fig. 14.1 13a-d], Fig. 11a, b [Fig. 14.114a, b]
Fig. lOa-c [Fig. 14.115a-d], Fig. 9a, c, d [Fig. 14.1 16a-c], Fig. 7a-c [Fig. 14.117a-c], Fig. 17a-c [Fig
14.1 18a-c], Fig. 16a, g [Fig. 14.1 19a-c], Fig. 14a-c [Fig. 14.121a-c], Fig. 15a-c [Fig. 14.122a-c], Fig
f,

22a, d, e [Fig. 14.124a-d], Fig. 21a, c, d [Fig. 14.126a-c], Fig. 19a, b, d [Fig. 14.128a-c], Fig. 20a, b
[Fig. 14.131a, b], Fig. 18a, e, f [Fig. 14.139a-c], Fig. 5a, b [Fig. 14.140a, b], Fig. 23a, g, h [Fig
14.142a-c], Fig. 25a-c [Fig. 14.144a-c], Fig. 26a-c [Fig. 14.145a-c], Fig. 28a, b [Fig. 14.147b, c], Fig
27a, g, h [Fig. 14.149a-c], Fig. 24a-c [Fig. 14.151a-c], Fig. 30a, b [Fig. 14.152a, b], Fig. 29a, d [Fig
14.156a, b], Fig. 63 [Fig. 14.163], Fig. 65 [Fig. 14.173], Fig. 58 [Fig. 14.174], Fig. 59 [Fig. 14.175], Fig
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With permission from Springer-Verlag GmbH & Co.:

Marchant, H.J. & Perrin, R. (1986), Planktonic choanoflagellates from two Antarctic lakes including the
descrition of Spiraloecion didymocostatum gen. et sp. nov., Polar Biol. 5: 207-210. Fig. 1 [Fig. 13.4a],
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Croome, R.L. (1987), Pinaciophora columna n. sp. and P. apora n. sp., new heliozoeans from Australia
and a report of P. fluviatilis Greeff from Antarctica, Arch. Protistenk. 131: 189-199. Fig. 7 [Fig. 15. 4d],
Fig. 10 [Fig. 15.5a]

Marchant, H.J. & Perrin, R. (1990), Seasonal variation in abundance and species composition of
choanoflagellates (Acanthoecidae) at Antarctic coastal sites, Polar Biol. 10: 499-505. Fig. 1 [Fig. 13.2a]

Valbonesi, A. & Luporini, P. (1990b), Description of two new species of Euplotes and Euplotes rariseta
from Antarctica, Polar Biol. 1 1 : 47-53. Fig. 6 [Fig. 14.29a]

Marchant, H.J., van den Hoff, J. & Burton, H.R. (1987b), Loricate choanoflagellates from Ellis Fjord,
Antarctica including the description of Acanthocorbis tintinnabulum sp. nov., Proc. NIPR Symp. Polar
Biol. 1: 10-22. Fig. 3 [Fig. 13.1c], Fig. 7 [Fig. 13.8a], Fig. 10 [Fig. 13.8c], Fig. 11 [Fig. 13. 8d], Fig. 14

[Fig. 13.1a]

Song, W. & Wilbert, N. (2000), Ciliates from Antarctic sea ice. Polar Biol. 23: 212-222. Fig. 2a, e
[Fig. 14.127a, b]. Fig. 3a, c [Fig. 14.159a-c], Fig. 4a, c [Fig, 14.150a, b]

With permission from Schweizerbart'sche Verlagsbuchhandlung:

Aescht, E. & Foissner, W. (1992), Biology of a high-rate activated sludge plant of a pharmaceutical
company, Arch. Hydrobiol., Suppl. 90: 207-251. Fig. 5 [Fig. 14.143a]

With permission from Servicio de Hidrografia Naval de la Armada Argentina:

Balech, E. (1962), Plancton de las campanas oceanograficas Drake I y II, Serv. Hidrogr. Naval Argent.
H. 627: 5-57. Fig. 14.50

With permission from Urban & Fischer Verlag:

Allgen, C.A. (1955), Die Suctorien der schwedischen Sudpolar-Expedition (1901-1903), Zool. Anz.
154: 36-48. Fig. 6 [Fig. 14.134]

Matthes, D. (1956), Suktorienstudien VIII. Thecacineta calix (Schroder 1907) (Thecacinetidae nov. fam.)
und ihre Fortpflanzung durch Vermoid-Schwarmer, Arch. Protistenk. 101: 477-528. Fig. 8 [Fig. 14.133]

Faure-Fremiet, E. & Ganier, M.-C. (1970), Structure fine du Strombidium sulcatum CI. et L. (Ciliata,
Oligotrichida), Protistologica 6: 207-223. Fig. 1 [Fig. 14.104a]

Fenchel, T. & Lee, C.C. (1972), Studies on ciliates associated with sea ice from Antarctica. I. The
nature of the fauna, Arch. Protistenk. 1 14: 231-236. Fig. lb [Fig. 14.125]

Grain, J., Puytorac, P. de & Groliere, C.A. (1982), Quelques precisions sur l'ultrastructure et la position
systematique du cilie Mesodinium rubrum, et sur la constitution de ses symbiontes chloroplastiques,
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Corliss, J.O. & Snyder, R.A. (1986), A preliminary description of several new ciliates from the
Antarctic, including Cohnilembus grassei n. sp., Protistologica 22: 39-46. Fig. 7 [Fig. 14.18b, c], Fig.
4 [Fig. 14.158]

Tamar, H. (1992), Four marine species of Mesodinium (Ciliophora: Mesodiniidae). II. Mesodinium
& Lachm., 1858, Arch. Protistenk. 141: 284-303. Fig. [Fig. 14.109]
pulex Clap. 1

Petz, W. (1994), Morphology and morphogenesis of Strombidium kryalis nov. spec. (Ciliophora,
Strombidiida) from Antarctic sea ice, Arch. Protistenk. 144: 185-195. Figs 1-3, 6 [Fig. 14.103a-c]

Petz, W. (1995), Morphology and morphogenesis of Thigmokeronopsis antarctica nov. spec, and T.

crystallis nov. spec. (Ciliophora, Hypotrichida) from Antarctic sea ice, Eur. J. Protistol. 31: 137-147.
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Song, W. & Packroff, G. (1997), Taxonomische Untersuchungen an marinen Ciliaten aus China mit
Beschreibungen von zwei neuen Arten, Strombidium globosaneum nov. spec, und S. platum nov. spec.
(Protozoa, Ciliophora), Arch. Protistenk. 147: 331-360. Fig. 17a [Fig. 14.15a], Fig. 15a, d, f [Fig.
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Agatha, S. & Riedel-Lorje, J.C. (1998), Morphology, infraciliature, and ecology of some strobilidiine
ciliates (Ciliophora, Oligotrichea) from coastal brackish water basins of Germany, Eur. J. Protistol. 34:
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Florentin, 1901, Strombidium sulcatum Claparede & Lachmann, 1859 and Heterostrombidium
paracalkinsi Lei, Xu & Song, 1999 (Ciliophora, Oligotrichida), Eur. J. Protistol. 36: 327-342. Fig. 18
[Fig. 14.104b], Fig. 19 [Fig. 14.104c]

With permission from Wiley-VCH:

Foissner, W. (1979), Okologische und systematische Studien iiber das Neuston alpiner Kleingewasser,
mit besonderer Berucksichtigung der Ciliaten, Int. Rev. Ges. Hydrobiol. 64: 99-140. Fig. 16 [Fig.
14.111a]

Illustrators, photographers and individual copyright-holders who provided images or

gave permission for them to be reproduced

*: Publications more than 50 years old, or the author deceased, or no contact possible following
repeated attempts.

Cover: V.Tucker.

Fig. 1.1: J. Cox.

Fig. 2.1: D.Thomas; 2.2a-c: F.Scott; 2.2d: G.Nash; 2.3a, d: F.Scott; 2.3b: L.Armand; 2.3c:
A.Ishikawa; 2.4: D.Thomas; 2.5: F.Scott; 2.6: D.Thomas; 2.7a: L.Armand; 2.7b: F.Scott; 2.7c, d:
D.Thomas; 2.8: F.Scott; 2.9: D.Thomas; 2.10: D.Thomas; 2.11: F.Scott; 2.12a, b, d: F.Scott; 2.12c:
K.Westwood; 2.13a: D.Thomas; 2.13b: K.Westwood; 2.13c: F.Scott; 2.13d: R.Perrin (from Perrin, Lu
& Marchant, 1987); 2.14: From Cupp, 1943*; 2.15a-d: D.Thomas; 2.16a, c-f: D.Thomas; 2.16b:
D.Thomas (from Everitt & Thomas, 1986); 2.17: D.Thomas; 2.18a-f, i: From Cupp, 1943*; 2.18g:
From Manguin, 1957*; 2.18h: From Hustedt, 1957*; 2.19-2.26: D.Thomas; 2.27a-c: D.Thomas; 2.27d,
e: F.Scott; 2.28-2.31: D.Thomas; 2.32a, b, d-f: D.Thomas; 2.32c: D.Thomas (from Everitt & Thomas,
1986); 2.33-2.35: D.Thomas; 2.36a, b, g, h: K.Westwood; 2.36c-f: F.Scott; 2.37a, e, f: L.Armand
(from Armand & Zielinski, 2001); 2.37b-d: L.Armand; 2.38a, b, d, e: D.Thomas; 2.38c: F.Scott; 2.39:
D.Thomas; 2.40a-c: F.Scott; 2.40d: E. van Wijk; 2.40e: S.Campbell; 2.41-2.42: D.Thomas; 2.43:
F.Scott; 2.44-2.56: D.Thomas; 2.57a, b: K.Westwood; 2.57c, d: D.Thomas; 2.58-2.65: D.Thomas;
2.66a, b, d: D.Thomas; 2.66c: F.Scott; 2.67a, b: D.Thomas; 2.67c: D.Thomas (from Everitt & Thomas,
1986); 2.67d: From Manguin, 1957*; 2.68a, c, d: F.Scott; 2.68b: D.Thomas; 2.69-2.70: F.Scott; 2.71:
D.Thomas; 2.72a-c: F.Scott; 2.72d: D.Thomas; 2.73a, c, d: F.Scott; 2.73b: J. van den Hoff; 2.73e:
Anna McEldowney; 2.74a, b: D.Thomas; 2.74c, d: F.Scott; 2.75a-c: D.Thomas; 2.75d, e: F.Scott;
2.76a, b: D.Thomas; 2.76c, d: F.Scott; 2.77-2.78: D.Thomas; 2.79a-c: F.Scott; 2.79d, e: D.Thomas;
2.79f, g: L.Cunningham; 2.80: D.Thomas; 2.81a-c: F.Scott; 2.81d, e: D.Thomas; 2.82a: K.Westwood;
2.82b: D.Thomas; 2.82c, d: F.Scott; 2.83a-c, e: F.Scott; 2.83d: D.Thomas; 2.84: D.Thomas; 2.85a, b:
D.Thomas; 2.85c-e: F.Scott; 2.86: D.Thomas; 2.87a: A.Ishikawa; 2.87b-d: D.Thomas; 2.87e, f:
F.Scott; 2.88a, b, d, e: F.Scott; 2.88c: D.Thomas; 2.89a, e: A.Davidson; 2.89b-d: F.Scott; 2.90a-c:
F.Scott; 2.90d: D.Thomas; 2.91: F.Scott; 2.92a, d, e: F.Scott; 2.92b, c: D.Thomas; 2.93: D.Thomas;
2.94a, c-e: F.Scott; 2.94b: D.Thomas; 2.95: D.Thomas; 2.96: F.Scott; 2.97a: F.Scott; 2.97b-d:
D.Thomas; 2.98-2.99: F.Scott; 2.100a: J. van den Hoff; 2.100b, c: D.Thomas; 2.100d, e: F.Scott;
2.101a-c: D.Thomas; 2.101d: A.Ishikawa; 2.101e, f: F.Scott; 2.102a-c: F.Scott; 2.102d, e: D.Thomas;
2.103: F.Scott; 2.104a, b: D.Thomas; 2.104c: A.Ishikawa; 2.104d: F.Scott; 2.105a, b: D.Thomas;
2.105c, d: A.Ishikawa; 2.106: D.Thomas; 2.107a: F.Scott; 2.107b, c: A.Ishikawa; 2.107d-f: D.Thomas;
2.108a: A.Ishikawa; 2.108b-f: D.Thomas; 2.109a, b: D.Thomas; 2.109c-e: F.Scott; 2.110: A.Ishikawa;
2.111a, c-e: F.Scott; 2.111b: D.Thomas; 2.112a: K.Westwood; 2.112b-e: D.Thomas; 2.113a-e:
F.Scott; 2.113f, g: A.Ishikawa; 2.114: F.Scott; 2.115a-c: A.Ishikawa; 2.115d-f: F.Scott.

Fig. 3.1: S.Campbell; 3.2-3.3: F.Scott; 3.4a: F.Scott; 3.4b-d: S.Campbell; 3.5a, b: F.Scott; 3.5c, d
S.Campbell; 3.6-3.7: F.Scott; 3.8a, b, d, e: F.Scott; 3.8c: J. van den Hoff; 3.9a, b, d, e: F.Scott; 3.9c
J.van den Hoff; 3.10a, b, d: F.Scott; 3.10c: J. van den Hoff; 3.11: F.Scott; 3.12a, b: F.Scott; 3.12c, d
S.Campbell; 3.13a: S.Campbell; 3.13b-d: A.McMinn; 3.14: S.Campbell; 3.15a: S.Campbell; 3.15c, d
F.Scott; 3.16a, c, d: F.Scott; 3.16b: S.Campbell; 3.17a, b: A.McMinn; 3.17c, d: S.Campbell; 3.18a, b
S.Campbell; 3.18c, d: F.Scott; 3.19a: S.Campbell; 3.19b-d: A.McMinn; 3.20: F.Scott; 3.21a, b, d

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S.Campbell; 3.21c: F.Scott; 3.22a, d: F.Scott; 3.22b, c: S.Campbell; 3.23a, c, d: S.Campbell; 3.23b:
A.McMinn; 3.24: A.McMinn; 3.25: S.Campbell.

Fig. 4.1a-c: F.Scott; 4.1d: G.Nash; 4.2a, b, e, f: F.Scott; 4.2c, d: S.Campbell.

Figs 5.1-5.2: A.Davidson; 5.3a, b: R. van den Enden; 5.3c, d: G.Nash; 5.4a: 0.Moestrup (from
Moestrup, 1979); 5.4b: H.Marchant; 5.5a-d: G.Nash; 5.5e: H.Marchant; 5.6: G.Nash. 5.7a: R. van den
Enden; 5.7b, c: F.Scott; 5.7d: G.Nash; 5.8: G.Nash; 5.9a, b: G.Nash. 5.9c, d: Anna McEldowney; 5.10:
a, b: Andrew McEldowney; 5.10c, d: G.Nash; 5.11a, b: G.Nash; 5.11c, d: R. van den Enden.

EMIDAS; 6.1c: Andrew McEldowney; 6.1d: S.Renaud, EMIDAS;

Fig. 6.1a: C.Findlay; 6.1b: J.Bollman,
6.2a: Andrew McEldowney; 6.2c: S.Renaud, EMIDAS; 6. 2d: M.Knappertsbusch,
C.Findlay; 6.2b:
EMIDAS; 6.3a: C.Findlay; 6.3b: Andrew McEldowney; 6.3c: C.Findlay; 6.3d: J.Bollman, EMIDAS;
6.4a: H.Thomsen (from Thomsen, 1981); 6.4b, b inset, d: H.Thomsen (from Thomsen, Buck, Coale,
Garrison, & Gowing, 1988); 6.4c: C.Findlay; 6.5a: C.Findlay; 6.5b, c: H.Thomsen (from Thomsen,
0stergaard, & Hansen, 1991); 6.5d: H.Thomsen (from Thomsen, 1980a); 6.5e: H.Thomsen (from
Thomsen, Buck, Coale, Garrison, & Gowing, 1988); 6.6a: C.Findlay; 6.6b, c: M.Knappertsbusch,
EMIDAS; 6.6d: C.Findlay; 6.7a: J.Young, EMIDAS; 6.7b, c: H.Marchant; 6.7d: C.Findlay; 6.8a, a
inset, c, d, e: H.Thomsen (from Thomsen, Buck, Coale, Garrison, & Gowing, 1988); 6.8b: H.Thomsen
(from Thomsen, 1981).

Fig. 7.1a: J. van den Hoff; 7.1b-f: N.Vors; 7.2a: D.Thomas; 7.2b: J. van den Hoff; 7.2c-e: N.Vors; 7.3:
N.Vers; 7.4a: Andrew McEldowney; 7.4b, c: D.Thomas; 7.4d: F.Scott; 7.5-7.6: Andrew McEldowney;
7.7a, c, d: F.Scott; 7.7b: G.Nash.

Fig. 8.1a, b: F.Scott; 8.1c-e:I.Moro (from Moro, La Rocca, Dalla Valle, Moschin, Negrisolo &
Andreoli, 2002); 8.2a-c: van den Hoff; 8.2d: S.Molloy; 8.2e: F.Scott; 8.3a: S.Molloy; 8.3b, c:
J.

Andrew McEldowney; 8.3d, e: N.Daugbjerg (from Daugbjerg, 2000); 8.4: H.Marchant (from Marchant,
Buck, Garrison & Thomsen, 1989).

Fig. 9.1a: F.Scott; 9.1b, c: A.Davidson.

Fig. 10.1a, b: G.Nash; 10.1c, d: Anna McEldowney.

Figs 11.1-11.2: F.Scott; 11.3: J. van den Hoff.

Fig. 12.1a: F.Scott. 12.1b: H.Marchant (from Marchant, van den Hoff & Burton, 1987).

Fig. 13.1a, c: J. van den Hoff (from Marchant, van den Hoff & Burton, 1987); 13.1b: F.Scott; 13. Id: J.

van den Hoff; 13.2a: H.Marchant (from Marchant & Perrin, 1990); 13.2b: H.Thomsen; 13.2c: Andrew
McEldowney; 13. 2d: H.Marchant; 13.3a, b, c: G.Nash; 13.3d: H.Marchant; 13.4a: G.Nash (from
Marchant & Perrin, 1986); 13.4b, c: H.Marchant; 13.4d: G.Nash; 13.5a, c, d: F.Scott; 13.5b:
H.Marchant; 13.5e: G.Nash; 13.6a, b, d: H.Marchant; 13.6c: H.Thomsen; 13.7a, b: G.Nash; 13.7c, d:
R. Perrin (from Marchant & Perrin, 1986); 13.8a, c, d: J. van den Hoff (from Marchant, van den Hoff &
Burton, 1987); 13.8b: H.Marchant; 13.9a: H.Marchant (from Marchant & Perrin, 1986); 13.9b:
H.Thomsen (from Thomsen & Boonruang, 1984); 13.9c. E.Takahashi (from Takahashi, 1981); 13. 9d:
H.Thomsen (from Thomsen, Buck, Coale, Garrison & Gowing, 1990); 13. 9e, f: B.Leadbeater (from
Leadbeater, 1972).

Fig. 14.1a: From Faure-Fremiet, 1936*; From

Ringuelet, 1953a*; 14.2: From Kahl, 1932*; 14.3:
14.1b:
M.Mulisch (from Mulisch, Harry, Patterson &
Wyatt, 1986); 14.4: From Faure-Fremiet, 1936*; 14.5-
14.6: From Kahl, 1932*; 14.7a,c: From Hadzi, 1951*; 14.7b: From Dons, 1914*; 14.8: From Ringuelet,
1957*; 14.9: From Faure-Fremiet, 1936*; 14.10: From Laackmann, 1910b*; 14.11: W.Song (from Song
& Wilbert, 2002); 14.12: W.Petz (from Petz,Song & Wilbert, 1995); 14.13a: From Kahl, 1932*;
14.13b: W.Song (from Song & Wilbert, 2002); 14.14: W.Petz (from Petz, Song & Wilbert, 1995);
14.15a: W.Song (from Song & Packroff, 1997); 14.15b, c: W.Petz (from Petz, Song & Wilbert, 1995);
14.16: W.Song (from Song & Wilbert, 2002); 14.17: W.Song (from Song & Packroff, 1997); 14.18a:
W.Petz (from Petz, Song & Wilbert, 1995); 14.18b, c: J.Corliss (from Corliss & Snyder, 1986); 14.19-
14.21: W.Song (from Song & Wilbert, 2002); 14.22: A.Valbonesi (from Valbonesi, 1996); 14.23-
14.25: W.Petz (from Petz, Song & Wilbert, 1995); 14.26: W.Song (from Song & Wilbert, 2002); 14.27:
W.Petz; 14.28: S.Serrano (from Serrano, Sola, Guinea, Arregui & Fernandez-Galiano, 1992); 14.29a-c:
W.Petz (from Petz, Song & Wilbert, 1995); 14.29d: A.Valbonesi (from Valbonesi & Luporini, 1990b);
14.30: S.Agatha (from Agatha, Spindler & Wilbert, 1993); 14.31: From Hada, 1970*; 14.32: From
Meunier, 1910*; 14.33: From Kofoid & Campbell, 1929*; 14.34a: From Balech, 1948*; 14.34b: From
Meunier, 1919*; 14.35: From Hada, 1970*; 14.36: From Meunier, 1919*; 14.37: From Meunier, 1910*;
14.38: From Hada, 1970*; 14.39: E. Balech (from Balech, 1971); 14.40a: From Laackmann, 1907*;
14.40b: V. Alder (from Alder, 1999); 14.41: W.Petz (from Petz, Song & Wilbert, 1995); 14.42a: From

547
 COPYRIGHT PERMISSION

Cleve, 1900*; 14.42b: From Brandt, 1906*; 14.43: From Cleve, 1900*; 14.44: From Hada, 1970*;
14.45a: W.Petz (from Petz, Song & Wilbert, 1995); 14.45b: From Laackmann, 1907*; 14.46a: From
Laackmann, 1907*; 14.46b: From Laackmann, 1910a*; 14.47: From Laackmann, 1907*; 14.48: From
Meunier, 1910*; 14.49: From Ostenfeld & Schmidt, 1902*; 14.50: E.Balech (from Balech, 1962b);
14.51-14.52: From Kofoid & Campbell, 1929*; 14.53: From Cleve, 1901*; 14.54a, b: From Small &
Lynn, 1985*; 14.54c: From Cleve, 1900*; 14.55: From Laackmann, 1910a*; 14.56: From Hada, 1970*;
14.57: From Brandt, 1906*; 14.58a-d: W.Petz (from Petz, Song & Wilbert, 1995); 14.58e: From
Kofoid & Campbell, 1929*; 14.59a-e: W.Petz (from Petz, Song & Wilbert, 1995); 14.59f: From
Laackmann, 1910a*; 14.59g, i: From Kofoid & Campbell, 1929*; 14.59H: From Laackmann, 1907*;
14.60: From Laackmann, 1910a*; 14.61a: From Laackmann, 1907*; 14.61b: From Laackmann, 1910a*;
14.62: From Laackmann, 1910a*; 14.63a: From Laackmann, 1907*; 14.63b: From Laackmann, 1910a*;
14.64a: From Laackmann, 1907*; 14.64b, c: From Laackmann, 1910a*; 14.65: From Hada, 1970*;
14.66: From Brandt, 1906*; 14.67: From Brandt, 1907*; 14.68: From Kofoid & Campbell 1929*;
14.69: From Laackmann, 1907*; 14.70: From Laackmann, 1910a*; 14.71-14.72: From Hada, 1970*;
14.73: From Brandt, 1906*; 14.74: From Hada, 1970*; 14.75-14.76: From Laackmann, 1910a*; 14.77:
From Jorgensen, 1924*; 14.78: W.Petz (from Petz, Song & Wilbert, 1995); 14.79: S.Agatha (from
Agatha & Riedel-Lorje, 1998); 14.80-14.81: W.Petz (from Petz, Song & Wilbert, 1995); 14.82:
D.H.Lynn (from Lynn & Montagnes, 1988); 14.83: W.Petz (from Petz, Song & Wilbert, 1995); 14.84:
From Thompson, 1972*; 14.85: W.Song (from Song & Wilbert, 2002); 14.86-14.88: W.Petz (from
Petz, Song & Wilbert, 1995); 14.89-14.90: W.Petz (from Petz, 1995); 14.91a: From Wulff, 1919*;
14.91b: D. J. S. Montagnes (from Montagnes, Lynn, Stoecker & Small, 1988); 14.92: W.Petz (from Petz,
Song & Wilbert, 1995); 14.93: D.H.Lynn (from Lynn, Montagnes & Small, 1988); 14.94: W.Petz
(from Petz, Song & Wilbert, 1995); 14.95: From Kahl, 1932*; 14.96: W.Petz (from Petz, Song &
Wilbert, 1995); 14.97-14.98: From Kahl, 1932*; 14.99: From Hada, 1970*; 14.100: D.H.Lynn (from
Lynn, Montagnes & Small, 1988); 14.101-14.102: W.Petz (from Petz, Song & Wilbert, 1995); 14.103:
W.Petz (from Petz, 1994); 14.104: W.Song (from Song, Wang & Warren, 2000); 14.105: W.Song (from
Song & Bradbury. 1998); 14.106a: From Wang, 1934*; 14.106b, c: W.Petz (from Petz, Song &
Wilbert, 1995); 14.107: W.Petz (from Petz, Song & Wilbert, 1995); 14.108a: T.Lindholm (from
Lindholm, 1985); 14.108b: J. Grain (from Grain, Puytorac & Groliere, 1982); 14.109a: H.Tamar (from
Tamar, 1992); 14.109b, c: From Borror, 1972*; 14.110: W.Petz (from Petz, Song & Wilbert, 1995);
14.111a: W.Foissner (from Foissner, 1979); 14.111b: J.Dragesco (from Dragesco & Dragesco-Kerneis,
1986); 14.111c: From Kahl, 1930*; 14.112: From Faure-Fremiet, 1924*; 14.113-14.119: W.Petz (from
Petz, Song & Wilbert, 1995); 14.120: W.Song (from Song & Wilbert, 2002); 14.121-14.122: W.Petz
(from Petz, Song & Wilbert, 1995); 14.123: W.Foissner (from Foissner & O'Donoghue, 1990); 14.124
W.Petz (from Petz, Song & Wilbert, 1995); 14.125: T.Fenchel (from Fenchel & Lee, 1972); 14.126
W.Petz (from Petz, Song & Wilbert, 1995); 14.127: W.Song (from Song & Wilbert, 2000a); 14.128
W.Petz (from Petz, Song & Wilbert, 1995); 14.129-14.130: W.Song (from Song & Wilbert, 2002);
14.131: W.Petz (from Petz, Song & Wilbert, 1995); 14.132: From Schroder, 1906*; 14.133: From
Matthes, 1956*; 14.134: From Allgen, 1955*; 14.135: From Allgen, 1952*; 14.136: From Schroder.
1911*; 14.137: From Schroder, 1906*; 14.138: W.Song (from Song & Wilbert, 2002); 14.139-14.140:
W.Petz (from Petz, Song & Wilbert, 1995); 14.141: From Hada, 1970*; 14.142: W.Petz (from Petz,
Song & Wilbert, 1995); 14.143a: E.Aescht (from Aescht & Foissner, 1992); 14.143b, c: W.Foissner
(from Foissner & Wilbert, 1981); 14.144-14.145: W.Petz (from Petz, Song & Wilbert. 1995); 14.146a:
From Maupas, 1883*; 14.146b: J.Dragesco (from Dragesco & Dragesco-Kerneis, 1986); 14.147a:
W.Song (from Song & Packroff, 1997); 14.147b, c: W.Petz (from Petz, Song & Wilbert, 1995); 14.148:
From Thompson, 1972*; 14.149: W.Petz (from Petz, Song & Wilbert, 1995); 14.150: W.Song (from
Song & Wilbert, 2000a); 14.151-14.152: W.Petz (from Petz, Song & Wilbert, 1995); 14.153: W.Song
(from Song, 2000); 14.154: W.Song (from Song & Wilbert, 2002); 14.155: W.Song (from Song, 2000);
14.156: W.Petz (from Petz, Song & Wilbert, 1995); 14.157: W.Song (from Song, 2000); 14.158:
J.Corliss (from Corliss & Snyder, 1986); 14.159: W.Song (from Song & Wilbert, 2000a); 14.160-
14.161: From Daday, 1913*; 14.162: W.Petz; 14.163: W.Petz (from Petz, Song & Wilbert, 1995);
14.164-14.167: W.Petz; 14.168: F.Scott; 14.169-14.172: W.Petz; 14.173-14.176: From Petz, Song &
Wilbert, 1995; 14.177-14.188: W.Petz; 14.189-14.193: F.Scott.

Fig. 15.1a, b: F.Scott; 15.1c, d: N.Vers; 15.2a, b: N.Vers; 15.2c:J. van den Hoff; 15.2d: F.Scott; 15.3:

Anna McEldowney; 15.4a, b, c: N.Vors; 15.4d: van den Hoff (from Croome, van den Hoff & Burton,
J.

1987); 15.5a: J. van den Hoff (from Croome, van den Hoff & Burton, 1987); 15.5b: N.Vers; 15.5c: J.
van den Hoff; 15.5d: R. Croome; 15.6a, c, d: N.Vors; 15.6b: A.Davidson; 15.7a: N.Vers; 15.7b:
F.Scott; 15.8a: N.Vors; 15.8b, c: F.Scott; 15.8d: A.Davidson.

548
 INDEX
Accepted names are in roman, synonyms and doubtful names in italic.

Acanthocorbis Aegyriana
apoda, 327 paroliva, 423
tintinnabulum. 326 Alexandrium
unguiculata, 327 tamarense, 222
Acanthocystidae, 456 Amastigomonas
Acanthocystis debruynei, 462
paliformis. 456 Amoeba
perpusilla, 457 spp., 449
turfacea, 457 Amoebae, 449
Acanthoeca Amphidinium
brevipoda, 327 acutissimum, 213
spectabilis, 327 hadai, 213
Acanthoecidae, 326 Amphipleura
Acanthoecopsis rut Hans, 148
asymmetrica, 328 var. antarctica. 148
unguiculata, 327 Amphiprora
Acanthostomella belgicae, 148
norvegica, 375 hyperborea, 134
Achnanthaceae, 121 var. genuina, 134
Achnanthes kjellmanii, 134
brevipes, 121 kufferathii, 133
vicentii, 122 paludosa
naviculacae, 122 f. m inula, 134

Acineta Amphisoleniacae, 206

calix, 427 Amphora
grahami, 427 angusta
ovoidea, 427 var. angustissima, 128
rotunda, 427 antarctica, 128
Acinetidae, 427 arcta, 128
Acropisthiidae, 415 bongrainii, 128
Actiniscus charcotii, 128
pennatus, 65 cymbelloides, 128
Actinocyclus gourdonii, 128
actinochilus, 52, 56 lanceolata, 128
adeliae, 52 var. aperta, 128
antarcticus, 110 var. robusta, 1 28

bifrons, 110 liouvellei, 128

cholnokyi, 55 mexicana
coscinodiscoides, 60 var. schmidtiana, 128
curvatulus, 52 peragallorum, 128
divisus, 55 var. robusta, 128
ehrenbergii, 55 petermannii, 128
excentricus, 90, 110 proteus, 128
exiguus, 55 var. oculata, 128
janus, 110 racovitzae, 128
octonarius, 55 var. minor, 128
oliverianus, 110 senonquei, 128
senarius, 51 turgida, 128
similis, 110 type A, 133
spiritus, 55 type B, 133
subocellatus, 52 type C, 133
umbonatus , 1 10 virgata, 128
undulatus, 51 var. crassa, 128
valdiviae, 110 Amphorella
Actinomonas antarctica, 380
mirabilis, 456 laackmanni, 387
Actinoptychus norvegica, 3 75
senarius, 51 Amphorellopsis
undulatus, 51 quinquealata, 387

549
 INDEX

Amphorides curvifilus. 461

laackmanni. 387 designis, 460
Ancyromonas marina, 461
melba, 462 saltans, 460
sigmoides, 462 Bodonida, 461
Anisonema Buehringa
prosgeobium, 319 antarctica, 400
Apusomonadidae, 463 Bursaopsis
Aristospira conicoides, 388
pachyderma, 452 ollula, 388
Ascampbelliellidae, 375 quinqaealata, 387
Ascobius Caecitellus
simplex, 348 parvulus, 462
irregularis, 348 Cafeteria
Aspidisca roenbergensis, 461
antarctica,359 Cafeteriaceae, 461
crenata, 359 Calciarcus
polypoda, 360 cf. alaskensis, 292, 294
quadrilineata, 360 Calcidiscaceae, 280
terranovae, 360 Calcidiscus, 280
Aspidiscidae, 359 leptoporus, 276, 277, 280
Asterolampra subsp. leptoporus, 280, 292
areolata, 14 subsp. quadriperforatus, 280, 293
heplactis, 13 Calciodinellaceae, 227
ralfsiana, 13 Calciopappus. 288
roperiana, 17 Calliacantha
Asterolampraceae, 13 natans, 333
Asteromphalus simplex, 333
buchii, 14 Calycomonas
cuvieri, 14 gracilis, 452
heptactis, 13 Calyptrosphaera
hookeri, 14 oblonga, 289, 294
humboldtii, 14 pirus, 289, 293
hyalinus, 14 Campanoeca
parvulus, 17 dilatata, 343
roperianus, 17 pedicellata, 337
Aulographonium Caneosphaera, 288
anthoides, 453 mollischii, 288
Auricula Cannopilus
compacta, 164 calyptra, 254
Auriculaceae, 164 Cannosphaera
Azpeitia antarctica, 453
neocrenulata, 55 Centrales, 13
nodulifera, 56 Centrohelida, 456
tabularis, 56 Ceratiaceae, 221
Bacillariaceae, 170 Ceratium, 221
Banquisia extensum, 221
belgicae, 148 fusus, 221
Berkeleya horridum, 202
rutilans, 148 kofoidii, 202
Bicosoecales, 461 lineatum, 221
Bicosta pentagonum, 221
antennigera, 328 subsp. grande, 222
minor, 328 petersii, 202
spinifera, 328 tripos, 222
Biddulphia Ceratoneis
arctica, 18 closterium, 170
aurita, 48 longissima, 191
litigiosa,48 Cercaria
polymorpha, 48 tripos, 222
striata, 48 Chaenea
weissjlogii, 48 elongata, 416
Blepharocysta teres. 416
paulsenii, 202 Chaetoceraceae. 21
splendor-maris, 231 Chaetoceros
Bodo subg. Chaetoceros, 21
cephaloporus, 462 subg. Hyalochaetae, 21

550
 INDEX

subg. Phaeoceros, 21 polylepis. 262

adelianus, 21 type A, 267
aequatorialis, 36 type B, 267
var. antarcticus. 21 type C, 267
atlanticum type D, 268
f. bulbosum, 27 type E, 268
atlanticus, 22 type F, 268
audax, 22 type G, 268
borealis, 35 type H, 268
brevis, 22 type 268
I,

brightwellii, 28 type 275

J,

bulbosus, 27 type K, 275

f. schimperiana, 27 type L, 275
castracanei, 27 type M, 275
chilensis, 36 vexillifera, 262
concavicornis, 28 Ciliophrydae, 456
convolutus, 28 Ciliophryida, 456
criophilus, 28 Claustrofolliculina
curvatus, 31 patagonica, 352
curvisetus, 31 Coccolithophorales, 276
debilis, 31 Coccolithus
dichaeta, 32 pelagicus, 276, 279
dispar, 22 Cocconeis
dislans, 32 costata
flexuosus, 32 var. antarctica, 122
gaussii, 32 var. pacifica, 127
gracile, 36 curiosa, 127
hendeyi, 35 fasciolata, 127
janischianum, 32 gautieri, 127
laciniosus, 35 imperatrix, 127
neglectus, 35 f. plena, 127
neogracilis, 36 infirmata, 127
pendulus, 21, 22, 36 litigiosa, 127
peruvianus, 28, 36 pacifica, 127
polygonum, 22 pell uc ida, 127
radicans, 37 pinnata
radiculum, 27 var. plena, 127
remotus, 32 schuettii, 127
schimperianus, 27 var. litigiosa, 127
septentrionalis, 36 var. minor, 127
simplex, 37 type A, 128
socialis, 37 Coccosphaera
tortissimus, 37 carteri, 281
Challengeron leptopora, 280
swirei, 453 Coccudina
Charcotia polypoda, 360
actinochilus, 52 Codonella
Chilodonella, 348 gaussi, 370
Chilomonas glacialis, 370
marina, 461 morchella, 370
Chlamydodontida, 420 naviculaefera, 373
Chlamydomonaceae, 316 prolongata, 373
Chlamydonella pusilla, 373
prostomata, 423 Codonellidae, 366
pseudochilodon, 423 Codonellopsidae, 369
Chlorodendrales, 308 Codonellopsis
Choanoflagellida, 326 antarctica, 369
Choreotrichida, 389 balechi, 369
Chroococcales, 324 frigida, 369
Chrysochromulina gaussi, 370
acantha, 260 glacialis, 370
bergenensis, 261 morchella, 370
chiton, 261 pusilla, 373
ericina, 261 soyai, 373
hirta, 261 turgescens, 370
leadbeateri, 262 Codonosigidae, 326
mantoniae, 268

551
 INDEX

Coelechinus var. typicus, 44

wapiticornis, 453 oppositus, 44
Cohnilembus pentas, 47
grassei, 439 pseudodenticulata, 89
Colepidae, 431 quadrifarius 121 ,

Coleps, 348 quinquemarcatus, 114

Conchacineta radiatus, 44, 56
rotunda, 427 var. asteromphalus, 38
Condylostoma var. oculus-iridis, 44
caudatum, 355 ritscheri, 113
granulosum, 355 sol, 84
remanei, 355 subtilis, 47, 114
Condylostomatidae. 355 symbolophorus, 47
Conferva var.? signatus, 47
rut Hans, 148 tabularis, 56
Conocylis var. antarctica, 56
helix. 399 trigonus, 114
Corbicula tumidus, 114
socialis, 339 \ar.fasciculatus, 114
Corethron Coscinosira
criophilum. 65 antarctica, 90
f. inerme, 60 oestrupii, 108
hispidum, 65 stellaris
inerme, 60 var. symbolophora, 47
murrayamtm, 65 Cosmoeca
pelagicum, 65 norvegica, 333
pennatum, 65 ventricosa, 334
valdiviae, 65 Cothurnia
Corynophrya antarctica, 445
campanula, 428 subglobosa, 445
interrupta, 428 Cothurniopsis
Coscinodiscaceae, 38 antarctica, 445
Coscinodiscus subglobosa, 445
actinochilus, 52 Coxliella
adumbratus 47 , cymatiocoides, 380
asteromphalus. 38 frigida, 374, 384
var. centralis, 38 intermedia, 374, 383
var. conspicua, 38 minor, 373
var. genuina, 38 Crinolina
bouvet, 38 aperta, 334
bullatus, 107 Cryothecomonas
centralis, 38 armigera, 463
chunii, 114 Cryptochilidae, 439
curvatulus, 43 Cryptochilum
var. divisus, 55 elegans, 435
var. subocellatus, 52 reniforme, 439
divisus, 55 Cryptomonadaceae, 317
eccentricus, 95 Cryptomonadales, 317
furcatus, 47 Cryptomonas
gracilis, 99 cryophila, 317
horridus, 43 Cryptophyceae, 477
incurvus, 114 Cryptophyte sp., 317
inflatus, 114 Crystallolithus
labyrinthus, 95 rigidus, 280. 292
lentiginosus, 100 Cyclidiidae, 445
leptopus, 107 Cyclidium
var. discrepans, 107 varibonneti, 445
lineatus, 107 Cyclotrichida. 408
margaritae, 107 Cyclotrichium
marginatus, 43 gigas, 411
minimus, 99 meunieri. 411
nodulifer, 56 Cylindrotheca
var. apiculata, 56 closterium. 170, 191
oculoides, 44 Cymatocylis
oculus-iridis, 38, 44 affinis, 379
var. genuina, 44 affmis/convallaria, 380
var. tenuistriata, 43 antarctica, 380

552
 INDEX

brevicaudata, 379 Diaphanoeca

calyciformis, 379 aperta, 334
calyx, 383 grandis, 337
conica, 383 multiannulata, 337
convallaria, 374, 379 pedicellata, 337
crassa, 380 sphaerica, 338
cristallina, 380 Diatoma
cu leu 1 1 us, 380 48
aurita,
cylindroides, 379 rhombicum, 183
cylindrus, 380 Dicladia
digitulus, 383 bulbosa, 27
drygalskii, 380 Dictyocha
ecaudata, 383 antarctica, 254
flava, 383 fibula, 251
f. typica, 380 octonaria, 254
folliculus, 383 speculum, 254
gaussi, 380 var. pentagonus, 254
glans, 380 var. septanarius, 254
kerguelensis, 383 Dictyochaceae, 251
nobilis, 383 Dictyochales, 251
ovata, 383 Dictyocysta
parva, 380 cassis, 379
scyphus, 380 Didiniidae, 411
subconica, 380 Didinium
subrotundata, 384 balbianii, 412
typica, 383 var. nanum, 412
vanhoeffeni, 383 gargantua, 411
Cymatosira nasutum, 412
laevis, 143 Dileptus, 348
Cymbellaceae, 128 Dinophysales, 206
Cytharoides Dinophysis, 206
australis, 360 antarctica, 206
balechi, 360 brevisulcus, 210
Cyttarocylididae, 379 contracta, 206
Cyttarocylis cornuta, 209
acus, 384 subsp. cornuta, 209
amor, 376 subsp. inerme, 209
calyciformis, 379 hastata, 206
cassis, 379 meteori, 209
conica, 379 norvegica, 209
drygalskii, 380 operculoides, 209
frigida, 374 ovum, 210
intermedia, 374 punctata, 210
minor, 373 rotundata, 213
nobilis, 383 simplex, 202
parva, 380 tenuivelata, 210
simplex, 376 tuberculata, 213
striata whittingae, 213
f. 376
curta, Diophrys
undella, 375 appendiculata, 356
Dactyliosolen marina, 356
antarcticus, 69 oligothrix, 356
f. typica, 69 scutum, 356
var. typica, 69 Diploneis
blavyanus, 66 stigmosa, 171
mediterraneus, 66 Diplopeltopsis
var. tenuis, 66 lenticula
meleagris, 66 f. minor, 228

tenuijunctus, 71 minor, 228

tenuis, 66 perlata, 228
Dadayiella Diplophrys
ganymedes, 388 archeri, 461
Daturella Diplopsalis
frigida, 388 lenticula, 228
Debya, 51 Discosphaera
Deutschlandia, 288 tubifera, 277

553
 77

INDEX

Dissodinium Euplotida, 356

lenticulum, 228 Euplotidae, 361
Distephanus Eupodiscaceae, 48
speculum, 254 Eupodiscus
Dubridinium tesselatus, 60
caperatum, 228 Eutreptiaceae, 319
Dysteria Eutreptiales, 319
calkinsi, 424 Eutreptiella
lanceolata, 424 cf. braarudii, 319
monostyla, 424 sp.,322
Dysteriida, 423 Exogenida, 424
Dysteriidae, 423 Exuviaella
Emiliania, 283 aequatoralis, 205
huxleyi, 276, 277, 283, 373 antarctica, 205
Enchelyodon baltica, 205
trepida, 412 compressa, 205
Endogenida, 427 elongata, 205
Entomoneidaceae, 133 lenticulata, 205
Entomoneis oblongata, 205
kjellmanii perforata, 203
var. kjellmanii, 134 Filellum
var. subtilissima, 134 expansum, 351
var. typica, 134 Folliculina
paludosa aculeata, 351
var. hyperborea, 134 ampulla, 348,351
Ephelota, 348 expansa, 351
Ephemera gigantea, 351
planamembranacea, 148 melitta, 355
Epiclintes,348 similis, 352
Epiplocylididae, 375 simplex, 348
Epiplocylis telesto, 352
mira, 375 violacea, 352
undella, 375 Folliculinidae, 348
Epiplocyloides Foraminiferida, 452
antarctica, 375 Fragilaria
Ervilia antarctica, 179
monostyla, 424 i.lata, 179
Eucampia brevistriata, 138
antarctica curta, 171
f. recta, 18 cylindrus, 172
f. spiralis, 1 f. elongata, 185
var. antarctica, 17 islandica, 137
var. recta, 18 var. adeliae, 137
balaustium, 1 7 var. islandica, 137
f. aestiva, 17 var. producta, 137
f. hiberna, 1 var. stricta, 137
var. minor, 17 linearis, 179
Eufolliculina nana, 172, 180
ampulacea, 348 obliquecostata, 1 79
Euglyphida, 452 f. maxima, 179
Euodia rhombica, 183
cuneiformis, 59 sublinearis, 180, 184
inornata, 59 f. longa, 184
radiata, 59 var. A,184
Euphysetta var.ambigua, 183
spp., 453 vanheurckii, 185
Euplotes Fragilariaceae, 137
acanthodus, 361 Fragilariopsis
algivorus, 366 antarctica, 179
antarcticus, 362 curta, 171
balteatus, 365 cylindrus, 172
euryhalinus, 365 f. minor, 172
focardii, 365 var. planctonica, 172
monostylus, 424 kerguelensis, 179
nohilii, 366 linearis, 179
rariseta, 366 var. curta, 171
sigmolateralis, 366 var. intermedia, 172

554
 INDEX

nana, 1 72 Gymnozoum
obliquecostata, 179 glaciale, 420
pseudonana, 180 sympagicum, 420
rhombica, 183 viviparum, 420
ritscheri, 183 Gyrodinium, 214
separanda, 184 glaciale, 217
sublinearis, 180, 184 lachryma, 217
vanheurckii, 185 rhabdomante, 217
Frontonia Gyrosigma
frigida, 432 sp., 153

Frontoniidae, 432 Halosphaeraceae, 308

Fuscheria Halteria
marina, 415 pulex, 408
Gaarderia, 288 rubra, 411
Gallionella Haptorida, 411
occulus, 66 Haslea
sol, 66 trompii, 153
Gastrocirrhidae, 360 Helicopontosphaera
Geminigera kamptneri, 282
cryophila, 311, 317 Helicosphaera, 281
Gephyrocapsa, 284 carteri, 293
caribbeanica, 284 var. carteri, 281
ericsonii, 284 kamptneri, 282
muellerae, 277, 284 Helicosphaeraceae, 281
oceanica, 284 Heliopeltaceae, 51
Gessnerium Hemiaulus
tamarensis, 222 antarcticus, 18
Glenodinium Hemidiscaceae, 51
acuminatum, 227 Hemidiscus
lenticula, 228 cuneiformis, 59
troichoideum, 227 kanayanus, 60
Globigerina Heterophrydae. 457
borealis, 453 Heterophrys
bu I hides myriapoda, 457
var. borealis, 452 Heteroschisma
pachyderma, 453 subantarctica, 213
Globorotalia Heterostentor
pachyderma, 453 coeruleus, 355
Globorotaliidae, 452 Heterotrichida, 348
Gomphonema Holosticha
kamtschaticum, 163 diademata, 395
Goniodomataceae, 222 foissneri, 395
Gonyaulacaceae, 226 pullaster, 395
Gonyaulacales, 221 spindleri, 395
Gonyaulax Hymenomonas
digitale, 226 roseola, 276
excavata, 222 Kakoeca
globosa, 222 antarctica, 338
scrippsae, 226 Katodinium
striata, 227 sp., 217
tamarensis, 222 Kentrophyllum
trygvii, 222 antarcticum, 416
Grammatophora Kinetoplastida, 461
arcuata, 138 Kolkwitziellaceae, 228
Gromia Kolpoda
oviformis, 452 lamella, 419
Guinardia Kryoprorodon
cylindrus, 73 garrisoni, 420
flaccida, 73 Kryoprorodontidae, 420
tubiformis, 73 Laackmanniella
Gymnodiniaceae, 213 naviculaefera, 373
Gymnodiniales, 213 prolongata, 373
Gymnodinium, 214 Laboea
baccatum, 214 conica
frigidum, 214 var. elongatum, 403
modestum, 214 crassula, 402
soyai, 214 emergens, 403

555
 7

INDEX

strobila, 399 Mesodinium

Lacrymaria pulex, 408
lagenula, 415 rubrum, 411
spiralis, 415 Mesoporos
Lacrymariidae, 415 perforatus, 203
Lagotia Metacylididae, 374
expansa, 351 Metacystis, 348
faurefremieti, 351 Metanophrys
gigantea, 351 antarctica. 440
similis. 352 Metaurostyla
simplex, 348 thompsoni, 396
Lauderia Metaurostylopsis
annulata. 84 rubra, 396
borealis, 84 Metromonas
glacial is, 84 simplex, 463
mediterranea, 66 Micropodiscus
Leegaardiella oliveriana, 110
elbraechteri. 390 Ministeria
Leegaardiellidae, 390 maricola, 462
Lembus Moelleria
pusillus, 432 antarctica, 1
Leprotintinnus Monodinium
gauss i, 370 balbianii,412
glacialis, 370 nanum, 412
naviculaeferus, 373 Monosiga
prolongatus, 373 ovata, 326
Leptocylindraceae, 60 Myrionecta
Leptocylindrus rubra, 347, 411
danicus, 65 Navicula
mediterraneus, 66 aspera, 164
Leucocryptos var. genuina, 164
marina, 461 challengeri, 154
Litonotidae, 416 criophila, 154
Litonotus cristata, 157
antarcticus. 416 crucicula, 158
emmerichi, 419 cryophila, 154
kopimorphus, 419 var. stauroptera, 154
lamella, 419 directa
Lohmanniella var. directa, 157
oviformis, 390 var. javanica, 157
spiralis, 389 gelida
Lohmanniellidae, 390 var. antarctica, 158
Loxophyllum var. parvula, 157
rostratum, 419 glaciei, 158
Luticola gracilis
muticopsis, 153 var. antarctica. 158
Lynchellidae, 423 kryophila, 154
Mamiellaceae, 312 mutica
Mamiellales, 312 var.cymbelloides 153 ,

Manguinea var. truncata,153

fusiformis, 154 muticopsiformis, 153
Mantoniella muticopsis, 153
antarctica, 312 f. capitata, 153
squamata, 312 f. evoluta, 153
Melosira f. reducta, 153
adeliae. 66 var. capitata, 153
antarctica, 69 pinnularia
dougetii. 90 var. constricta, 158
occulus, 66 planamembranacea, 148
radiala, 66 schefterae, 158
sol, 66 stauroneiformis, 164
Melosiraceae, 66 subpolaris, 157
Membraneis triplex, 157
challengeri, 154 trompii, 153
Meringosphaera var. major, 153
mediterranea, 307 Naviculaceae. 148
Mesodiniidae, 408

556
 INDEX

Nematacineta turgidula, 198

ovoidea, 427 turgiduloides, 198
rotunda, 427 vanheurckii, 185
Nematoplata Nitzschiella
bicuneata, 192 tenuirostris, 1 70
Neogloboquadrina Noelaerhabdaceae, 283
pachyderma, 452 Notocephalus
Nitzschia parvulus, 396
aciculariformis, 143 Ochromonadales, 295
acicularis, 185 Odontella
f. angustior, 188 aurita, 48
var. antarctica, 188 litigiosa, 48
var. interrupta, 188 weissflogii, 48
amphiprora, 187 Ophryodendridae, 424
angulata, 183 Ophryodendron
antarctica, 183 conicum, 424
australis, 187 Orchitophryidae, 440
barbieri. 187 Orthodonella
var. latestriata, 192 shenae, 428
barkleyi, 197 Orthodonellidae, 428
bicapitata, 188 Oxytoxaceae, 227
bilobata, 187 Oxytoxum
closteriodes, 198 criophilum, 227
closterium, 1 70 Pappomonas. 287
compacta, 164 weddellensis, 287
curta, 171 Papposphaera, 287
cylindrus, 172 arctica, 287, 293
decipiens, 188 borealis, 287, 293
delicatissima, 193 obpyramidalis, 287
dissipata polybotrys, 288, 293
var. antarctica, 188 sagittifera, 287
dubia, 187 sarion, 287
var. australis, 187 simplicissima, 288
frigida, 192 Papposphaeraceae, 287
heimii, 193 Paractedoctema
hybrida, 187, 191 acruosa, 445
hybridaeformis, 191 Parafolliculina
kerguelensis, 179 amphora, 352
lanceolata patagonica, 352
var. pygmaea violacea, 352
f. australis, 188 Paralia
var. spicula, 188 sol, 69
lecointei, 188 Parapetalotricha
lineata, 179 meridiana, 379
lineola, 197 Paraphysomonadaceae, 295
longissima, 191 Paraphysomonas
medioconstricta, 170, 191 antarctica, 295
obliquecostata, 180 butcheri, 295
ostenfeldii circumforaminifera, 301
var. minor, 187 cribosa, 296
pacific a, 193 cylicophora, 296
peragallii, 192 foraminifera, 301
prolongata, 197 imperforata, 296
prolongatoides, 197 oligocycla, 296
pseudohybrida, 191 type A, 301
pseudonana, 180 vestita, 301
183
ritscheri, Parauronema
separanda, 184 antarcticum, 435
sicula virginianum, 435
var. bicuneata, 192 Paraurostyla, 348
var. migrans, 192 Parmales, 301
stellata, 192 Parundella, 348
subcurvata, 198 Parvicorbicula
subhybrida, 191 circularis, 338
sublineata, 184 ongulensis, 339
taenia, 192 quadricostata, 339
taeniiformis 192 , socialis, 339

557
 INDEX

Paulinella Phalacroma, 206

intermedia, 452 contractum, 206
Paulinellidae, 452 cornuta, 209
Pedifolliculina operculoides, 209
arctica, 355 rotundatum, 213
Pelagacineta ruudii, 206
campanula. 428 spinata, 209
interrupta, 428 Philasterida, 432
Pelagostrobilidium Philasteridae, 439
neptuni, 389 Phyllomonas
spirale, 389 simplex, 463
Peniculida, 432 Physomonas
Pennales, 13, 121 vest it a, 301
Pentalamina Pinaciophora
corona, 305 denticulata. 460
Pentalaminaceae. 305 fluviatilis, 460
Peridinales, 227 Pinnularia
Peridinium aspera, 164
adeliense, 233 criophila, 154
affine, 233 direct a, 157
antarcticum, 235 quadratarea
applanatum, 236 var. constricta, 158
archiovatum, 236 var. soederlundii, 163

areolatum, 236 Placidae, 431

bellulum, 236 Placus
charcoti, 239 antarcticus, 431
concavum, 239 Plagiotropus
conicum australis, 163

f. islandica, 248 gaussii, 163

crassipes, 240 Planktoniella
cruciferum, 240 sol, 84
curtum, 240 Pleurasiga
defectum, 240 cupula, 345
depressum, 235 minima, 340
elegantissimum, 243 orculaeformis, 340
granii, 239, 243 reynoldsii, 340
incertum, 244 sphyrelata, 343
incognitum, 244 Pleurasiga
latistriatum, 244 complexa, 345
mediocre, 247 Pleurochloridaceae, 307
melo, 247 Pleuronema
metananum, 247 coronatum, 440
meunierii, 228 glaciale, 440
mite, 239 puytoraci, 440
obovatum, 236 Pleuronematida, 440
ovatum, 240, 247 Pleuronematidae, 440
penitum, 247 Pleurosigma
petersii, 239 directum, 163
rosaceum, 248 smithianum, 163
splendor-maris, 231 Pleurostomatida, 416
subinerme, 244 Ploesconia
sympholis, 248 balteata, 365
thulesense, 248 scutum, 356
troichoideum, 227 Podolampaceae, 231
unipes, 248 Podolampas
Peritromus, 348 antarctica, 231
Petalotrichidae, 379 Podosira
Petasaria adeliae, 89
468
heterolepis, 84
glacialis,

Phaeocystaceae, 255 hormoides

Phaeocystis var. glacialis, 84
globosa, 256 liotardii,47
pouchetii, 256 oliveriana, 110
Phaeocystis Polarella
antarctica, 255 glacialis. 218
Phaeodina Polyfibula
antarctica, 453 caudata, 343

558
 INDEX

elatensis, 343 Protoperidinium, 231

sphyrelata. 343 acanthophorum, 202
Polykrikaceae, 218 adeliense, 233
Polykrikos affine, 233
schwartzii, 218 antarcticum, 235
Polytoma applanatum, 236
papillata, 316 archiovatum, 236
Pontosphaera areolatum, 236
huxleyi, 283 bellulum, 236
Porella bipatens, 239
adriatica, 203 cf. crassipes, 240

asymmetrica, 203 charcoti, 239

bisimpressa, 203 concavum, 239
globulus, 203 cruciferum, 240
perforata, 203 curtum, 240
Porosira defectum, 240
anlarctica, 84 digitale, 226
glacialis, 84 elegantissimum, 243
pseudodenticulata, 89 glyptopterum, 243
Porotheca granii, 243
perforata, 203 incertum, 244
Porpostoma incognitum, 244
grassei, 439 joergensenii, 202
notatum, 439 latistriatum, 244
Postgaardi macrapiticum, 202
mariagerensis, 323 manginii, 202
Potamodiscus mediocre, 247
kalbei, 460 melo, 247
Preperidinium metananum, 247
granulosum, 202 pallidum, 202
meunierii, 228, 236 parcum, 202
perlatum, 228 parvicollum, 202
Proboscia, 73 penitum, 247
alata, 73 pseudoantarcticum, 202
f. inermis, 74 pyriforme, 202
inermis, 74 pyrum, 202
truncata,74 radius, 202
Prodinophysis raphanum, 202
contracta, 206 rosaceum, 248
rotundata, 213 saltans, 202
ruudii, 206 sphaeroideum, 202
Proplectella thulesense, 248
claparedii, 384 turbinatum, 202
globosa, 387 unipes, 248
Prorocentraceae, 203 variegatum, 202
Prorocentrales, 203 Protorhabdonella
Prorocentrum curta, 376
antarcticum, 205 simplex, 376
aporum, 205 Prymnesiaceae, 260
balticum, 205 Prymnesiales, 255
bidens, 205 Pseudobodo
compressum, 205 sp.,461
lebourae, 205 tremulans, 461
pomoideum, 205 Pseudocohnilembidae, 432
Prorodontida, 431 Pseudocohnilembus
Protocymatocylis longisetus, 432
pseudoconica, 384 marinus, 432
subrotundata, 384 persalinus, 432
Protocystis pusillus, 432
acornis, 453 Pseudofolliculina
micropelecus, 453 melitta, 355
triangularis, 453 Pseudogomphonema
tridens, 453 groenlandicum, 164
Protogonyaulax kamtschaticum, 163
excavata, 222 var. antarctica, 164
Protoperidiniaceae, 231 Pseudokeronopsidae, 396

559
 INDEX

Pseudomicrosportella curvata, 78
ornata, 327 cylindrus, 73
Pseudonitzschia hebetata
antarctica, 193 f. bidens, 77
australis,187 f. semispina, 81
barkleyi, 197 imbricata, 81
var. obtusa, 198 var. shrubsolei, 81
delicatissima, 193, 201 var. striata, 81
heimii, 193 inermis, 74
lineola, 197 f. castracanei, 74
prolongatoides, 197 f. rostrata, 74
sicula murrayana, 77
var. bicuneata, 192 obtusa, 73, 74
subcurvata, 198 polydactyla
turgidula, 198 f. polydactyla, 82
turgiduloides, 198 squamosa, 82
f.

Pseudostaurosira rhombus, 82
brevistriata, 138 rostrata, 74
Pseudotrachelocerca semispina, 81
trepida, 412 shrubsolei, 81
Pseudotrachelocercidae, 412 sima
Pteridomonas f. silicea, 83
danica, 456 f. sima, 82
Pterosperma simplex, 83
cf. polygonum, 308 striata, 81
parallelum, 308 styliformis, 83
polygonum, 308 f. latissima, 83
Ptychocylididae, 376 var. latissima, 82
Ptychocylis var. longispina, 82, 83
amor, 376 var. oceanica, 82
ostenfeldi, 376 var. polydactyla, 83
undella, 3 75 var. semispina, 81
vanhoeffeni, 383 tenuijuncta, 71
wailesi, 376 torpedo, 83
Pyramimonas truncata, 74
australis, 311 tubiformis, 73
gelidicola, 311 Rhizosoleniaceae, 69
tychotreta, 312 Rhynchomonas
Pyrodinium nasuta, 460
phoenus, 222 sp., 460
Pyxidicula Richoda
compressa, 205 pullaster, 395
Raphoneis Rimostrombidium
bicuneata, 192 conicum, 390
Rhabdoaskenasia, 348 glacicolum, 390
Rhabdonella Roperia
amor, 376 tesselata, 60
Rhabdonellidae, 376 var. coscinodiscoides, 60
Rhaphoneis Rotosphaerida, 457
fasciolata, 127 Saepicula
Rhizomonas leadbeateri, 343
setigera, 66 Salpingella
Rhizosolenia, 77 acuminatoides, 388
acuminata, 77 aff. 388
faurei,
alala, 73, 74 costata, 389
f. curvirostris, 74 decurtata, 389
f. genuina, 73 laackmanni, 388
f. indica, 74 Salpingoeca
f. truncata, 74 natans, 333
antarctica, 73 spinifera, 328
antennata Schimperiella
f. antennata. 77 antarctica, 1 10
f. semispina, 77 oliveriana, 1 10
bidens, 77 valdiviae, 110
chunii, 77 Scrippsiella
crassa, 78 faeroense, 227
curva, 78 troichoidea. 227

560
 INDEX

Scyphidia, 348 pseudocinctum, 399

Semifolliculina pulchrum, 400
gigantea, 351 rhyticollare, 400
similis, 352 striatum, 407
simplex, 348 strobilum, 399
Sessilida, 445 sulcatum, 404
Spatangidium syowaensis, 404
heptactis, 13 tintinnodes, 404
ralfsianum, 13 wulffi, 407
Sphenomonadaceae, 319 Stylonychia
Sphenomonadales, 319 appendiculata, 356
Spiniferites Styloplotes
bulloideus, 227 appendiculatus, 356
Spiraloecion Suessiaceae, 218
didymocostatum, 344 Suessiales, 218
Spirodinium Symbolophora
lachryma, 217 furcata, 47
Spiroprorodon microtrias, 47
garrisoni, 420 Synechococcus
glacialis, 420 sp., 324
Spirostrombidium Synedra
pseudocinctum, 399 acicularis, 185
rhyticollare, 400 fragilis, 143
Sportelloeca nitzschioides, 144
kentii, 346 var. minor, 144
Stauroneis pelagic a, 147
achnanthes, 164 reinboldii, 147
aspera, 164 tabulata
Stauroptera var. thalassiothrix, 147
aspera, 164 thalassiothrix, 147
Stellarima Synedropsis
microtrias, 47 fragilis, 143
Stenosemella hyperboreoides, 143
avellana, 374 laevis, 143
oliva, 374 lata
perpusilla, 374 144
var. angustata,
Stentoridae, 355 var. lata,143
Stephanoeca recta, 137, 144
complexa, 345 Synhymeniida, 428
cupula, 345 Synurales, 307
kenti, 346 Syracolithus
norrisii, 346 catilliferus, 282, 293
Strobilidiidae, 389 confusus, 282, 293
Strobilidium quadriperforatus 280, 293
,

anlarcticum, 400 Syracosphaera, 288

conicum, 390 anthos, 277
diversum, 403 corrugis, 288
elegans, 405 elatensis,288
elongatum, 403 molischii, 288
neptuni, 389 pulchra, 276, 277, 289
spirale,389 oblonga type, 294
striatum, 407 pirus type, 293
sulcatum, 404 quadriperforata, 280, 293
syowaensis, 404 Syracosphaeraceae, 288
Strombidiida, 399 Tachysoma
Strombidiidae, 399 parvulum, 396
Strombidium Telonema
acutum, 400 antarctica, 468
antarcticum, 400 subtile, 468
conicoides, 401 Terebraria
crassulum, 402 kerguelensis, 179
diversum, 403 Tetraparma
elongatum, 403 pelagica, 302
emergens, 403
glaciale. 404 Thalassionema
hadai, 404 nitzschioides, 144
kryale, 404 f. curta, 144

561
 INDEX

Thalassiosira Tintinnida, 366

ambigua, 89 Tintinnidae, 387
antarctica, 89 Tintinnopsis
var. borealis, 89 avellana, 374
var. echinata, 147 bacillaria, 366
anliqua buetschlii, 366
var. septata, 108 glans, 369
australis, 90 lata, 367
delicatula, 99 lobiancoi, 367
dichotomica, 95 loricata, 366
eccentrica, 95 major, 366
expecta, 99 oliva, 374
fallax, 89 parva, 369
ferelineata, 96 petasa, 369
frenguellii, 96 Tintinnus
frenguelliopsis, 97 acuminatoides, 388
gerloffii.91,99 acuminatus
gracilis var. secata, 388
var. expecta, 99 claparedii, 384
var. gracilis, 99 costatus, 389
gravida, 100 ganymedes 388 ,

lentiginosa, 100 quinquealatus, 387

leptopus, 107 Tokophrya
maculata, 107 campanula, 428
margaritae, 107 interrupta, 428
oestrupii, 108 steueri, 428
oliveriana, 110 Tontonia
partheneia, 110 antarctica, 408
perpusilla, 111 Torodinium
poro-irregulata, 112 sp.,217
poroseriata, 113 Trachelius
ritscheri, 113 teres, 416
rotula, 121 Trachelocerca
scotia,113 trepida, 412
symmetrica, 114 Trachelophyllidae, 416
tcherniai,100 Trachyneis
114
trifulta, aspera, 164
tumida, 114 164
var. antarctica,
Thalassiosiraceae, 84 var.genuina, 164
Thalassiothrix Trachysphenia
antarctica, 147 australis
curvata, 144 var. kerguelensis, 179
frauenfeldii Triceratium
var. nitzschioides, 144 arcticum, 18
gelida, 144 Trichoda
longissima, 147 transfuga, 358
var. antarctica, 147 Trichotoxon
nitzschioides, 144 reinboldii, 147
Thaumatomastix Trigonaspis, 294
fragilis, 469 melvillea, 294
fusiformis, 469 Trigonium
splendida, 469 arcticum, 18
type A, 469 Triparma
Thecacineta columacea
calix, 427 subsp. alata, 302
oblonga, 427 laevis
paradesmodorae, 427 subsp. pinnatilobata, 302
subantarctica, 427 subsp. ramispina, 305
Thecacinetidae, 427 strigata, 305
Thecomonas Triparmaceae, 302
trahens, 462 Trochilia, 348
Thigmokeronopsis Tropidoneis
antarctica, 396 antarctica, 154
crystallis, 399 belgicae, 148
Tiarina fusiformis, 154
antarctica, 431 gauss ii, 163
fusus, 206

562
 INDEX

Turrisphaera transfuga, 358

arctica, 287, 293 uncinata, 359
borealis, 287, 293 Uropedalium
polybotrys, 288, 293 antarcticum, 436
Umbellosphaera, 290 Urostyla
irregularis, 290 sp., 396
Umbellosphaeraceae, 289 Urostylida, 395
Umbilicosphaera Urostylidae, 395
hulburtiana, 277 Vaginicolidae, 445
Undella Volvocales, 316
antarctica, 387 Vorticella
claparedii. 384 ampulla, 348
declivis, 387 Wigwamma, 292
hyalina, 387 annulifera, 292, 294
Undellidae, 384 antarctica, 292
Urocyclon arctica, 292
ovatum, 435 triradiata, 292
Uronema Woodania, 401
acutum, 435 Xystonella
antarcticum, 435 acus, 384
elegans, 435 Xystonellidae, 379
marinum, 432, 436 Zosterodasys
paramarinum, 436 kryophilus, 431
virginianum, 435 Zygabikodinium
Uronematidae, 435 lenticulatum, 228
Uronychia perlatum, 228
antarctica, 358 Zygoceros
binucleata, 359 caballeroi, 48
heinrothi, 359

563
 Protists are microscopic algae and

protozoa, formerly thought of as

single-celled plants and animals.

Planktonic protists constitute

the base of marine food webs £*'

and play a key role in global

ocean-atmospheric

gas exchange.

This book is a

comprehensive guide

to the protists that li 1

in the surface waters and

sea-ice around Antarctica.

More than 550 organisms

are described and superbly

illustrated with over

1300 light and electron

micrographs and drawings.

The inclusion of an

extensive bibliography of

the widely scattered literature

makes Antarctic Marine Protists

an indispensible resource for

marine biologists.

' Australia n Government

Department of the Environment and Heritage
BioYogica"
Resources Australian Biological Resources Study
Stad y Australian Antarctic Division
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