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MARINE ZOOPLANKTON
PRACTICAL GUIDE
for the Northwestern Arabian Gulf

Volume 1

Faiza Yousif Al-Yamani


Valeriy Skryabin
Aleksandra Gubanova
Sergey Khvorov
Irina Prusova
Kuwait Institute for Scientific Research, Kuwait

2011
MARINE ZOOPLANKTON PRACTICAL GUIDE
for the Northwestern Arabian Gulf

Published in Kuwait in 2011 by


Kuwait Institute for Scientific Research,
P.O. Box 24885, 13109 Safat, Kuwait

Copyright © Kuwait Institute for Scientific Research, 2011


All rights reserved

Design by
Mariposa Marketing & Advertising
Creative Director: Melad Helani

Printed and bound by


Lucky Press

First Edition
Kuwait Institute for Scientific Research (Publisher)
ISBN 978-99966-95-07-0

No part of this work may be reproduced or utilized in any form or by any means
electronic or manual, including photocopying, recording or by any information
or retrieval system, without the prior written permission of the Kuwait Institute
for Scientific Research.

Salmiya beach - Photo by Dr. V. Skryabin - Kuwait Institute for Scientific Research

M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Table of CONTENTS
PREFACE i

ACKNOWLEDEGEMENTS iii

INTRODUCTION 1

MATERIALS AND METHODS 2

1. PROTOZOA 8

1.1. Tintinnida 8

Glossary 9

Key to Families of the Order Tintinnida 11

List of the Arabian Gulf Species Encountered During 1999 – 2010 12

Identification and Taxonomy of Tintinnid Species from the Arabian Gulf 14

Family TINTINNIDIIDAE 14
Genus Leprotintinnus 14
Leprotintinnus bubiyanicus 14
Leprotintinnus elongatus 16
Leprotintinnus nordqvisti 18
Family CODNELLIDAE 20
Genus Tintinnopsis 20
Tintinnopsis acuminata 20
Tintinnopsis ampla 22
Tintinnopsis angusta 24
Tintinnopsis baltica 25
Tintinnopsis beroidea 26
Tintinnopsis compressa 27
Tintinnopsis dadayi 28
Tintinnopsis directa 30
Tintinnopsis failakkaensis 32
Tintinnopsis gracilis 34
Tintinnopsis karajacensis 36
Tintinnopsis lobiancoi 37
Tintinnopsis lohmannii 38
Tintinnopsis mortensenii 39
Tintinnopsis nana 40
Tintinnopsis orientalis 41
Tintinnopsis parva 42

K u wait I n s titute f or Scienti f ic R e s earc h


Tintinnopsis parvula 43
Tintinnopsis radix 44
Tintinnopsis rotundata 46
Tintinnopsis sacculus 47
Tintinnopsis schotti 48
Tintinnopsis tocantinensis 50
Tintinnopsis turbo 52
Tintinnopsis undella 53
Tintinnopsis urnula 54
Family CODONELLOPSIDAE 56
Genus Codonellopsis 56
Codonellopsis lusitanica 56
Codonellopsis morchella 58
Codonellopsis ostenfeldi 59
Family DICTYOCYSTIDAE 60
Genus Dyctiocysta 60
Dyctiocysta duplex 60
Genus Luminella 62
Luminella kuwaitensis 62
Family METACYLIDIDAE 64
Genus Coxliella 64
Coxliella annulata 64
Genus Helicostomella 66
Helicostomella longa 66
Genus Metacylis 68
Metacylis jorgenseni 68
Metacylis lucasensis 70
Metacylis pithos 72
Metacylis tropica 74
Metacylis sp. 75
Family ASCAMPBELLIELLIDAE 76
Genus Ascampbelliella 76
Ascampbelliella obscura 76
Family PETALOTRICHIDAE 78
Genus Petalotricha 78
Petalotricha ampulla 78
Family RHABDONELLIDAE 80
Genus Protorhabdonella 80
Protorhabdonella curta 80
Protorhabdonella simplex 81
Genus Rhabdonella 82
Rhabdonella conica 82

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Rhabdonella striata 84
Rhabdonella sp. 85
Family XYSTONELLIDAE 86
Genus Parundella 86
Parundella lohmanni 86
Genus Xystonella 87
Xystonella treforti 87
Genus Xystonellopsis 88
Xystonellopsis gaussi 88
Genus Favella 89
Favella adriatica 89
Favella campanula 90
Favella ehrenbergi 91
Favella panamensis 92
Family UNDELLIDAE 94
Genus Undella 94
Undella dilatata 94
Family TINTINNIDAE 96
Genus Amphorellopsis 96
Amphorellopsis acuta 96
Genus Amphorides 98
Amphorides amphora 98
Amphorides quadrilineata 98
Genus Dadayiella 100
Dadayiella cuspis 100
Dadayiella ganymedes 102
Genus Eutintinnus 103
Eutintinnus apertus 103
Eutintinnus conicus 104
Eutintinnus contractus 106
Eutintinnus fraknoi 108
Eutintinnus lusus-undae 109
Eutintinnus macilentus 110
Eutintinnus tenue 111
Eutintinnus turgescens 112
Eutintinnus sp. 113
Genus Salpingella 114
Salpingella attenuata 114
Salpingella rotundata 116
Genus Salpingacantha 117
Salpingacantha unguiculata 117
The Geographic Distribution of Tintinnid Species 118

K u wa i t I n s t i t u t e f o r S c i e n t i f i c R e s e a r c h
1.2. Zoothamnium 120
Zoothamnium sp. 120

1.3. Ellobiopsis 122


Ellobiopsis chattoni 122

1.4. Noctiluca 124
Noctiluca scintillans 124

2. METAZOA 126
2.1. Cnidaria 126
Amphinema rugosum 126
Aequorea pensilis 128
Eirene viridula 130
Eutima gegenbauri 132
Octophialucium funerarium 134
Obelia spp. 136
Podocoryne sp. 138
Clytia discoida 140
Solmundella bitentaculata 142
Cunina octonaria 144
Aglaura hemistoma 146
Liriope tetraphyla 148
Sanderia malayensis 150
Diphyes chamissonis 152

2.2. Ctenophora 154
Pleurobrachia pileus 154
Beroe sp. 156

2.3. Platyhelminthes 158


Acoelomorpha gen.spp. 158
Turbellaria Larvae 160

2.4. Nemertea 162


Pilidium Larvae 162

2.5. Rotifera 164


Families Synchaetidae and Dicranophoridae 164

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2.6. Annelida 166
Lopadorhynchus henseni 166
Pelagobia longicirrata 168

2.7. Brachiopoda 170


Brachiopod’s Larvae 170

2.8. Mollusca 172
Atlanta sp. 172
Limacina bulimoides 174
Creseis chierchiae 175
Pneumodermatidae gen.sp. 176
Sepiolidae gen.sp. 177

2.9. Lophotrochozoa Larvae 178


Actinotrocha Larvae 178
Cyphonautes Larvae 180

2.10. Cirripedia 182


Balanus amphitrite 182

2.11. Cladocera 184


Penilia avirostris 184
Pleopis polyphemoides 186
Pseudevadne tergestina 188

2.12. Ostarcoda 190


Euconchoecia aculeata 190
Cypridina sp. 192

Index of Scientific Names 194

K u wa i t I n s t i t u t e f o r S c i e n t i f i c R e s e a r c h
The marine zooplankton community of Kuwait
waters includes fascinating organisms, which are
abundant, diverse, and encompasses representatives
of all the major invertebrate phyla. Along with
PREFACE
phytoplankton, zooplankton are key components of
marine ecosystems forming the base of most marine
food webs. Plankton constitute the principal diet of
the early life stages of fish including commercially
important fish.

Zooplankton research is an essential component


in forming a more complete understanding of
the functioning of marine ecosystems. However,
identifying zooplankton can be a challenging and a
tedious task and requires a high level of expertise.

The samples used to produce this guide


were obtained during the 1999-2010 sea cruises
covering all Kuwait waters. A total of 183 species of
zooplankton were recorded and identified. Species
diversity was highest for the groups of tintinnids,
copepods, and cnidarians (75, 54 and 14 species,
respectively).

This illustrated zooplankton guide of Kuwait


waters is an invaluable reference for taxonomists,
marine plankton ecologists, environment managers,
coastal engineers, students of invertebrate biology,
environmental impact assessment experts and
marine biologist. This practical book provides a
comprehensive documentation of zooplankton
biodiversity and taxonomical description of the
commonly encountered species in Kuwait waters as
well as the Arabian Gulf.

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“We consider species to be like
a brick in the foundation of a
building. You can probably lose
one or two or a dozen bricks and
still have a standing house. But by
the time you’ve lost 20 per cent of
species, you’re going to destabilize
the entire structure. That’s the way
ecosystems work.”

Donald Falk

K u wa i t I n s t i t u t e f o r S c i e n t i f i c R e s e a r c h ii
Gratitude is extended to Mr. Alan Lennox (Kuwait
Institute for Scientific Research, Kuwait) for his
tremendous help in collecting the needed samples
for this study from Kuwait waters. Our appreciation
Acknowledgements
to Mrs. Gracekutty Thomas Vargese for her hard
work in mesozooplankton sample analysis, and to
Mr. Vasiliy Prusov for his help in computerizing the
Copepod drawings. We thank Mrs. Linda Fernandes
for compiling and formatting the materials used for
this guide.

The authors are grateful to the Kuwait Institute


for Scientific Research for the financial support of this
research project.

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We share this planet with many species. It is our responsibility to
protect them, both for their sakes and our own.

Pamela A. Matson

K u wa i t I n s t i t u t e f o r S c i e n t i f i c R e s e a r c h iv
Each species on our planet plays a role in the healthy functioning of
natural ecosystems, on which humans depend.

William H. Schlesinger

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Zooplankton are tiny drifting organisms living in
the water and drifting at the mercy of currents.
They are vital components of the marine food
Introduction chains. Plankton communities integrate various
human and environmental impacting factors, hence,
providing a benchmark for monitoring the synergistic
effects of urbanization and climate change (Kunz
and Richardson 2006; Richardson and Kunz 2006).
Long-term zooplankton data provide an important
baseline against which future changes in the marine
ecosystem can be assessed.
Zooplankton can be subdivided into
holoplankton, i.e., permanent members of
the plankton (e.g., calanoid copepods), and
meroplankton, i.e., temporary members in the
plankton (e.g., larvae of fish, shrimp, and crab). The
meroplankton group consists of larval and young
stages of animals that will adopt a different lifestyle
once they mature.
Zooplankton organisms are categorized by their
size into macro-, meso-, and micro-zooplankton
(Table 1). They range from bacteria to jellyfish
(microns to centimeters in size).
Zooplankton are the initial prey item for almost
all fish larvae as they switch from their yolk sacs to
external feeding as well as for many plankton-eating
adult fishes. Natural factors (e.g., current variations,
climate change, etc.) and anthropogenic factors (e.g.,
increasing pollution, dams on rivers, etc.) can impact
zooplankton, and hence, affect the survival of fish
larvae, and hence, the fisheries resources.
Zooplankton encompasses representatives of all major invertebrate phyla, including
some that can only be found in the plankton. Three guides have already been produced for
Kuwait zooplankton groups: for the tintinnids (Al-Yamani and Skryabin, 2006), the copepods
(Al-Yamani and Prusova, 2003) and larval decapods (Al-Yamani and Khovorov, 2010). However,
several additions and updates are now available to be added to the aforementioned guides on
the zooplankton of Kuwait waters. Hence, this guide will include all these updates, including
descriptions of several other zooplankton groups for Kuwait waters, as well as descriptions and
documentation of some major zooplankton groups for the Arabian Gulf waters.
This illustrated guide includes descriptions, drawings and microphotographs for all the
major zooplankton groups that were encountered in the samples. The guide will help users to
identify their specimens into major taxa and to species level.

Table 1. Size Categories of Zooplankton

Term Size Examples


Macrozooplankton > 20 mm (> 2 cm) Metazoans: Chaetognaths; Medusae; Ctenophores; Salps, Doliolids, adult and
juvenile Decapods
Mesozooplankton 0.2-20 mm Metazoans: Copepods; Cladocerans; Ostracods; Chaetognaths; Tunicates;
Pteropods; Heteropods
Microzooplankton 20-200 µm Protozoans: Foraminiferans; Radiolarians; Ciliates.
Metazoans: Rotifers; juvenile stages of marine organisms (e.g., Copepod nauplii)

K u wait I n s titute f or Scienti f ic R e s earc h 1


The sampling area in Kuwait waters (the north-
western corner of the Arabian Gulf ) extended from
Khor Al-Sabbiyah and Bubiyan waters in the north
Materials and Methods to the southern waters of Kuwait toward the Saudi
Arabian border south of Al-Khiran. The distance of
the sampling stations from the coastline varied from
less than 1 km to approximately 45 km offshore.
Monthly sampling for Kuwait waters was
conducted from 1999-2010 covering a total
of 47 stations (Figs. 1a, 1b and 1c). Additional
microzooplankton samples were examined, which
were collected from 41 stations in the Arabian Gulf
during a regional cruise organized by the Regional
Organization for the Protection of the Marine
Environment, extending from Kuwait’s waters to the
Strait of Hormuz waters (Fig. 1d). Stations’ depths
varied from 5 to 95 m.
The microzooplankton samples were collected
from Kuwait waters as well as regional waters using
a 20 µm mesh net, with 20 cm mouth diameter,
by vertical or oblique tows. The plankton net was
towed for 1-5 minutes depending on the density of
plankton at each sampling location. Samples were
placed into one-liter plastic bottles and immediately
fixed with Lugol preservative (40 ml /L).
Before processing, the sample was mixed
carefully and a subsample was taken with a
calibrated Stempel-pipette. The subsample was
examined under a dissecting stereomicroscope
with magnification of 20x35 using a Bogorov’s or
Utermöhl’s counting chambers. If the sample was
rich in microzooplankton, a 4 ml subsample was
examined, however, if the abundance of organisms
was low, three to five subsamples of 4 ml each
were processed. Microzooplankton organisms in
the whole chamber were identified to the lowest
taxon possible. Some samples contained detritus,
inorganic particles or phytoplankton, which masked
the observation of small organisms, such as the
tintinnids with transparent hyaline lorica. In that
case, one drop of cytoplasm staining Rose Bengal
was added into the subsample to make these
organisms more visible.

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“Man cannot discover new oceans unless he has the courage to
lose sight of the shore.”

Andre Gide

K u wait I n s titute f or Scienti f ic R e s earc h 3


Fig. 1 a. Zooplankton sampling stations in Kuwait waters (July 2002 – June 2005).

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Fig. 1 b. Zooplankton sampling stations in Kuwait waters (1999 – 2010).

K u wait I n s titute f or Scienti f ic R e s earc h 5


Fig. 1 c. Stations sampled off Bubiyan Island (February 2004 – February 2005).

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Fig. 1 d. Location of ROPME microzooplankton sampling stations in the Arabian Gulf and the Strait of Hormuz
(February – March 2006).

For proper identification, measurement, and microphotography of microzooplankters such


as tintinnids, the specimens were placed on a glass slide, immersed in a drop of glycerol to avoid
fast drying of the objects to be observed, and examined under the compound microscope with
magnification of 10x10 – 10x100. Microzooplankton organisms were picked up from the counting
tray by the use of a micropipette.
The mesozooplankton samples were collected from Kuwait waters by oblique tows using a 110
µm mesh net, with mouth opening of 50 cm diameter. The duration of the tows ranged from 1-5
minutes depending on the abundance of plankton at the sampling location. Samples were carefully
transferred into one-liter plastic bottles and immediately preserved with 5% formaldehyde solution,
which was previously buffered with disodiumtetraborate (borax, Na2B4O7.10 H2O).
The mesozooplankton samples were processed using the method of portions (Yashnov, 1939).
Samples were reduced to volumes of 150, 200 or 250 ml depending upon zooplankton density, and
mixed to ensure that all organisms were distributed randomly in the sample volume. Subsequently,
a 1 ml, or 5 ml, or 10 ml subsample (depending on the volume of zooplankton content) was taken
by a calibrated Stempel-pipette, and all the specimens less or equal to 1 mm in total length were
identified and enumerated. Large (above 1 mm body length) and rare specimens were identified
and enumerated in 1/2, or 1/4, or 1/8, or 1/16, or 1/32 portion of the sample. To split the sample
into portions, the Motoda Plankton Splitter (Motoda, 1959) was used. Identification and size
measurements of organisms were made in the Bogorov’s counting chamber under the dissecting
stereomicroscope. All organisms were identified to the lowest taxon possible.

K u wait I n s titute f or Scienti f ic R e s earc h 7


The dissection of copepods was carried out on the glass 1. PROTOZOA
slides under dissecting stereomicroscope with fine needles.
Two dissecting needles were used. With one needle the 1.1. Tintinnida
specimen was held firmly and with the other needle the
somites, legs or mouthparts of the copepod were cut. Kingdom Alveolata
For identification, drawing and microphotographing, the Phylum Ciliophora Doflein, 1901
mesozooplankton specimens and dissected parts were Subphylum Intramacronucleata Lynn, 1996
placed on glass slides, cleared in glycerol and observed Class Spirotrichea Bütschli, 1889
under the compound stereomicroscope with magnification Subclass Choretrichia Small and Lynn, 1985
of 10x10 – 10x100. Order Tintinnida Kofoid and Campbell, 1929
A thorough literature search was conducted for the
identification of the different groups of zooplankton Kingdom Alveolata (groupings of organisms that share
that were encountered in Kuwait waters. For protozoan many characteristics): Alveolates are protists with sub-
microzooplankton, a general reference book chapter on surface cavities (alveoli) and complex internal structures.
ciliated protozoa by Lynn and Small (2000) was used, as They include both free-living and parasitic forms. Three
well as references on tintinnids of Kofoid and Campbell major subgroups are recognized: the ciliates, distinguished
(1929), Hada (1932, 1937), Marshall (1969), Paulmier (1997), by the presence of two nuclei and cilia; the dinoflagellates,
Fernandes (2004) and Al-Yamani and Skryabin (2006). A mostly unicellular with nuclei and with two flagella; and the
very useful site “The World of Protozoa, Rotifera, Nematoda apicomplexans, a group of parasites.
and Oligochaeta” of Inamori (1996-2010) [http://web2.nies.
go.jp/chiiki1/protoz] was utilized for the description of Phylum Ciliophora: Cell body covered with cilia,
some of the protozoan species included in this guide. nulcei differentiate to macro- and micronuclei, cilia also
The identification of mesozooplankton, particularly, of differentiate to unique structures (cirri, membranelles, etc),
copepods was carried out with the assistance of relevant consisting of many species.
references: Al-Yamani and Prusova (2003), Bradford et al.
(1983), Bradford-Grieve (1994, 1999), Bradford-Grieve et Subphylum Intramacronucleata: Microtubules associated
al. (1999), Brodsky et al. (1983), Mori (1937), Rose (1933), with the division of the macronucleus, and are located
Silas and Pillai (1973) and Vervoort (1963). The Internet within the macronucleus.
site “Diversity and Geographic Distribution of Marine
Planktonic Copepods” of Dr. C. Razouls et al. (2005-2010) Class Spirotrichea: Intramacronucleate ciliates may
[http://copepodes.obs-banyuls.fr/en] was also used for have somatic dikinetids usually with the anterior or both
the description of some copepod species included in this kinetosomes ciliated or somatic polykinetids; generally,
guide. Several zooplankton references, such as Michel et with conspicuous right and left oral and/or preoral ciliature,
al. (1986a and 1986b); Conway et al. (2003) and Marine with left serial oral polykinetids leading (usually clockwise)
Species Identification Portal (2000-2010) [http://species- into the oral cavity, either around a broad anterior end or
identification.org/index.php], were used to identify the along anterior and left margins of the body; polykinetids
other groups of zooplankton. Taxonomic position of (undulating membranes) may also be present along the
organisms was validated using the Internet site of Appeltans right side of the oral region; the cytostome is deep in
et al. (2000-2010) World Register of Marine Species [http:// some groups and shallow in others; the body is ciliated
www.marinespecies.org]. in some groups, but nearly devoid of cilia in others; some
are loricate, others are not; the lorica may be attached to
substrate or not.

Subclass Choreotrichia: Spirotrich ciliates, body generally


conical or bell-shaped; body cilia poorly developed; serial
oral polykinetids form closed, outer circle around broader
anterior end; the inner ends of some of these outer
polykinetids may extend into the oral cavity where they
may accompany a smaller number of oral polykinetids
restricted to the inner oral cavity; most are planktonic.

Order Tintinnida: Choreotrich ciliates, with lorica, attached


by aboral process to inner wall of lorica.

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Glossary

Ciliates of order Tintinnida produce lorica which is a hard


or soft case for protection. It can be transparent or covered
with foreign particles. Ciliates are fastened to lorica with an
offshoot of the cell called pedicel. Tintinnids move in the
water using membranelles, each of them is formed with
some connected cilia. Tintinnids can leave their loricas and
later build new ones, which usually differ from the primary
loricas.
L - hyaline lorica of family Xystonellidae;
OE - oral end of lorica;
AE - aboral end of lorica;
M - membranelles;
Ma - macronucleus;
Mi - micronucleus;
P - pedicel;
AH - aboral horn

Lorica of tintinnids of the family Codonellopsidae has


transparent hyaline collar, often with spiral structures and
opaque agglomerated bowl.
C - collar;
B - bowl;
F - fenestrae

Tintinnids of the genus Luminella (family Dictyocystidae)


have short hyaline collar with squarish or semi-circular
windows.

Tintinnids of the genus Eutintinnus (family Tintinnidae)


have transparent hyaline lorica with open aboral end.
OE - oral end of lorica;
AE - aboral end of lorica.

K u wait I n s titute f or Scienti f ic R e s earc h 9


Tintinnids of the genus Tintinnopsis (family Codonellidae)
have agglomerated loricas of various shapes.
OR- oral rim;
C - collar;
B - bowl.

Some tintinnids during their life time add to their loricas


one or more secondary oral rims.
P - primary oral rim;
S - secondary oral rim.

Some tintinnids have wide oral expansion called oral flare


(e.g. Tintinnopsis mortensenii).
OF - oral flare.

All tintinnids of the genus Leprotintinnus (family


Tintinnidiidae) have open aboral end, often with aboral
flare.
AF - aboral flare.

Basic measurements of lorica (family Metacylididae):


OD - oral diameter;
MT - maximum transdiameter;
CL - length of collar;
BL - length of bowl;
TL - total length of lorica.

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Key to Families of the Order Tintinnida

1. Lorica with foreign material, either agglomerated or agglutinated, lightly or densely covered, which is possibly restricted

in distribution on or within lorica wall ……………………………………………………….……………………………………………………………………………………….…… 2

1´. Lorica not as above …………………………………………………………….……………………………………………………………………………………….……………………………… 5

2. Lorica collar with one or two rows of oval or quadrangular fenestrae ………………………………………………….......……………Dictyocystidae

2´. If collared, without fenestrae ……………………………………………………….………………………………………………………………………………….……………......……… 3

3. Lorica wall soft and gelatinous ……………………………………………………….…………………………………………......……………………….…………Tintinnidiidae

3´ Lorica wall firm ……………………………………………………….………………………………………………………………………………………….…………………………….....………… 4

4. Lorica covered with agglomerated particles …………………………………………………….……………………………………………………...….… Codonellidae

4´. With hyaline collar, no or sparse agglutination on collar ………………………………………………….…………………………………Codonellopsidae

5. Lorica hyaline, with rings or spiral bands on part or entire lorica ……………………………………………….…………………....……… Metacylididae

5´. Lorica not as above ……………………………………………………….……………………………………………………………………………………….…….........………………………… 6

6. Lorica hyaline, short, tubular or cup-shaped; with two collars, outer may be toothed…………………..………… Ascampbelliellidae

6´. Lorica not as above ……………………………………………………….…………………………………………………………………………………….........….……………………………… 7

7. Lorica hyaline, with flared lip on top of a vertical cone; small suboral fenestrae on lip and bowl ………………………………………………
……….……………………………………………………………………………………….……………………………………………………………………….......…………….…… Petalotrichidae

7´. Lorica not as above ……………………………………………………….………………………………………………………………………………….………....……………………………… 8

8. Lorica hyaline, with longitudinal, low ribs that may be simple, branched or anastomosed ……………………..…… Rhabdonellidae

8´. Lorica not as above ……………………………………………………….……………………………………………………………………………………………....…………………… ……… 9

9. Lorica hyaline, usually smooth, chalice-shaped, with horn; meshwork between wall layers has inconspicuous, thin wall

material separating polygons ……………………………………………………….……………………………………………………………….…….......………… Xystonellidae

9´. Lorica not as above ……………………………………………………….……………………………………………………………………………………….…………………………………… 10

10. Lorica hyaline, goblet-shaped; inner and outer layers of lorica wall conspicuously thick ……………………………………………………… Undellidae

10´. Lorica not as above ……………………………………………………….……………………………………………………………………………………….………………………………… 11

11. Lorica hyaline, smooth, without rings, bands or ribs; inner and outer layers of wall thin …………………………………………...………… Tintinnidae

K u wait I n s titute f or Scienti f ic R e s earc h 11


List of the Arabian Gulf Species Encountered During 1999 - 2010

Phylum Class Subclass Order Family Genus / Species

Ciliophora Spirotrichea Choreotrichia Tintinnida Tintinnidiidae Leprotintinnus


Leprotintinnus bubiyanicus
Leprotintinnus elongatus
Leprotintinnus nordqvisti
Codonellidae Tintinnopsis
Tintinnopsis acuminata
Tintinnopsis ampla
Tintinnopsis angusta
Tintinnopsis baltica
Tintinnopsis beroidea
Tintinnopsis compressa
Tintinnopsis dadayi
Tintinnopsis directa
Tintinnopsis failakkaensis
Tintinnopsis gracilis
Tintinnopsis karajacensis
Tintinnopsis lobiancoi
Tintinnopsis lohmannii
Tintinnopsis mortensenii
Tintinnopsis nana
Tintinnopsis orientalis
Tintinnopsis parva
Tintinnopsis parvula
Tintinnopsis radix
Tintinnopsis rotundata
Tintinnopsis sacculus
Tintinnopsis schotti
Tintinnopsis tocantinensis
Tintinnopsis turbo
Tintinnopsis undella
Tintinnopsis urnula
Tintinnopsis spp.
Stenosemella sp.
Codonellopsidae Codonellopsis
Codonellopsis lusitanica
Codonellopsis morchella
Codonellopsis ostenfeldi
Dictyocystidae Dyctiocysta
Dyctiocysta duplex
Luminella
Luminella kuwaitensis
Metacylididae Coxliella
Coxliella annulata
Helicostomella
Helicostomella longa
Metacylis
Metacylis jorgenseni
Metacylis lucasensis
Metacylis pithos
Metacylis tropica
Metacylis sp.

12 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Phylum Class Subclass Order Family Genus / Species

Ascampbelliellidae Ascampbelliella
Ascampbelliella obscura
Petalotrichidae Petalotricha
Petalotricha ampulla
Rhabdonellidae Protorhabdonella
Protorhabdonella curta
Protorhabdonella simplex
Rhabdonella
Rhabdonella conica
Rhabdonella striata
Rhabdonella sp.
Xystonellidae Parundella
Parundella lohmanni
Xystonella
Xystonella treforti
Xystonellopsis
Xystonellopsis gaussi
Favella
Favella adriatica
Favella campanula
Favella ehrenbergi
Favella panamensis
Undellidae Undella
Undella dilatata
Tintinnidae Amphorellopsis
Amphorellopsis acuta
Amphorides
Amphorides amphora
Amphorides quadrilineata
Dadayiella
Dadayiella cuspis
Dadayiella ganymedes
Eutintinnus
Eutintinnus apertus
Eutintinnus conicus
Eutintinnus contractus
Eutintinnus fraknoi
Eutintinnus lusus-undae
Eutintinnus macilentus
Eutintinnus tenue
Eutintinnus turgescens
Eutintinnus sp.
Salpingella
Salpingella attenuata
Salpingella rotundata
Salpingacantha
Salpingacantha unguiculata

K u wait I n s titute f or Scienti f ic R e s earc h 13


Identification and Taxonomy of Tintinnid Species from the Arabian Gulf

Family: TINTINNIDIIDAE Kofoid and Campbell, 1929


Reference: Lynn and Small, 2000.

With agglomerate on soft, tubular lorica; lorica may be attached to the substrate; no collar; most are
marine, some freshwater; two genera.

Genus Leprotintinnus Jörgensen, 1899

References: Kofoid and Campbell, 1929; Hada, 1937


Tintinnidiidae with lorica elongated, tube-like, open at both ends; frequently, but not always with
spiral structures as in Coxliella either for the whole or part of the length of the lorica; no collar;
surface viscous, freely agglomerating foreign bodies; wall soft and coarsely alveolar. Differs from
Tintinnidium in the widely open aboral end.
The genus includes 5 described species:
Leprotintinnus bottnicus (Nordqvist) Jorgensen
Leprotintinnus neriticus (Campbell)
Leprotintinnus nordqvisti (Brandt) (Brandt) Kofoid and Campbell
Leprotintinnus pellucidus (Cleve) Jorgensen
Leprotintinnus simplex Schmidt
In Kuwait waters three species were found, including two new species, namely, Leprotintinnus
elongatus and Leprotintinnus bubiyanicus, in addition to Leprotintinnus nordqvisti. Description of
these three species of Leprotintinnus from Kuwait waters are included below.

Leprotintinnus bubiyanicus Skryabin and Al-Yamani, 2007

References: Al-Yamani and Skryabin, 2006; Skryabin and Al-Yamani, 2007 (Fig. 2 a-f ).

Description: Lorica soft and fragile, cylindrical, never with flaring oral and aboral ends, sometimes
slightly narrowing toward the aboral end, up to 3 degrees (Fig. 2 a, b, c). Wall often shows a weak
spiral structure, more pronounced at aboral half of lorica, it usually depends both on the shape of
the wall and of the distribution of particles (Fig. 2 d); agglomeration usually is relatively poor and
even at the oral part of lorica and more dense at the aboral part.

Measurements of Specimens from Kuwait Waters: Total length 471.7 µm (350.2-834.3 µm).
Average oral diameter 72.8 µm (72.1-82.4 µm). Average aboral diameter 70.7 µm (61.8-72.1 µm).
Approximate ratio L/oral diameter 4.9-10.1. Thus, it is the longest species of all known tintinnids.
Lorica of L. bubiyanicus, as well as lorica of L. elongatus, is very fragile, especially in the aboral part,
such that the minimum size of this tintinnid is not identifiable. Usually there are many fragments of
loricas in the samples (loricas could be broken during sampling or transportation), which makes it
difficult to determine the real size of tintinnids.

Comments: L. bubiyanicus differs from L. nordqvisti in the absence of the aboral funnel and longer
lorica, from L. bottnicus and L. pellucidus in the absence of sharp narrowing of the aboral end, from
L. simplex in larger size, from L. elongatus in the absence of oral rim and from L. neriticus in smaller
width, larger length, and spiral structure formed not only by distribution of particles, but also by the
shape of the wall.

Distribution: Leprotintinnus bubiyanicus is abundant in the northern waters of Kuwait off Boubiyan
Island but only in winter and early spring; its maximum abundance reached 150 X 103 organisms/m3
in February 2004 (Skryabin and Al-Yamani, 2007a). In spring this tintinnid produces one (Fig. 2 e) or
two (Fig. 2 f ) cysts and survives the warm period as a resting stage.

14 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


b

e
f

Fig. 2. Leprotintinnus bubiyanicus. Scale – 50 µm.


a-b. loricas of different length and shape; c. the longest of observed tintinnids, the length of this specimen is 834.3
µm; d. structure of lorica; e. L. bubiyanicus with one cyst; f. L. bubiyanicus with two cysts.

K u wait I n s titute f or Scienti f ic R e s earc h 15


Leprotintinnus elongatus Skryabin and Al-Yamani, 2007
References: Al-Yamani and Skryabin, 2006; Skryabin and Al-Yamani, 2007 (Fig. 3 a-d).

Description: Lorica almost cylindrical, with slightly flaring oral and aboral ends (Fig. 3 a-b), long
specimen usually slightly narrowing toward the aboral end (up to 4 degrees). Long specimens often
have one (Fig. 3 c) or two (Fig. 3 d) secondary oral rims. Wall showing a faint spiral structure, lorica
agglomerated more thickly in the aboral end. Most of the long specimens in the samples have
damaged aboral end without rim.

Measurements of Specimens from Kuwait Waters: Total length 245.9 µm (212.5-332.5 µm);
average oral diameter 37.0 µm (32.5-42.5 µm). Approximate ratio of L/oral diameter 5.3-9.5. Lorica of
this tintinnid is very fragile, especially in the aboral part; most of the tintinnids in the samples were
more or less damaged, which made it difficult to measure exactly the minimum size of this tintinnid
species.

Comments: L. elongatus differs from L. nordqvisti in the absence of the aboral funnel, from L.
bottnicus, and L. pellucidus in the absence of sharp narrowing of the aboral end, from L. simplex in
flaring oral end and spiral structure of the wall and from L. neriticus (380-470 µm) by its smaller size
(212.5-332.5 µm).

Distribution: Leprotintinnus elongatus was observed at all sampled stations, but occurred in low
numbers during all seasons, however, its abundance was higher (up to 83 X 103 organisms/m3) in
winter.

16 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

Fig. 3. Leprotintinnus elongatus. Scale – 50 µm.


a, b. loricas without secondary oral rim; c. lorica with one secondary oral rim; d. loricas with two secondary oral rims.
Arrows indicate Secondary oral rims.

K u wait I n s titute f or Scienti f ic R e s earc h 17


Leprotintinnus nordqvisti (Brandt, 1906)
(Fig. 4 a-f )

Description: Lorica consisting of a tubular shaft and an inverted funnel-shaped aboral flare, 4.3-10.5
oral diameters in length; oral rim irregular, usually slightly flaring (80-85 degrees); shaft more or less
tapering (15 degrees), expanding near the posterior region to form a distinct aboral conical flare (50-
110 degrees); aboral aperture large, 1.2-2.3 oral diameters; aboral margin very ragged; wall showing
a faint spiral structure, made of rather scare particles aggregated more thickly on the surface of the
aboral flare than on that of the shaft.
The species differs from the other species of Leprotintinnus in having a conspicuously large aboral
flare.

Reported Measurements: Length 245 µm (150-352 µm); oral diameter 35 µm (30-38 µm); aboral
diameter 61 µm (48-80 µm).

Measurements of Specimens from Kuwait Waters: Most of the Arabian Gulf specimens of L.
nordqvisti differ by size and, especially by proportions. They are shorter and wider, often with large
aboral flare. Total length 159±3 µm (128-192 µm); oral diameter 43±1 µm (109-154 µm); aboral flare
129±1 µm (109-154 µm). Some specimens of L. nordqvisti have secondary oral rims (Fig. 4 e).

Distribution: Leprotintinnus nordqvisti is common in Kuwait waters and around Bubiyan Island. This
species occurred along the Iranian coast with southeastern direction extending to 28°N and 51°30´E
and was not observed in the central, eastern and southern regions of the Gulf. Abundance of L.
nordqvisti was higher in winter and early spring.

18 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

e f

Fig. 4. Leprotintinnus nordqvisti. Scale – 50 µm.


a - c. shape of lorica of L. nordqvisti typical for Kuwait waters; d. shape of lorica more typical for other regions of the
World Ocean; e. lorica with secondary oral rims; f. aboral flare.

K u wait I n s titute f or Scienti f ic R e s earc h 19


FAMILY: CODNELLIDAE Kent, 1881
Reference: Lynn and Small, 2000

With agglomerate on flask-, bowl- or chalice-shaped lorica; collar not clear, if present may or may not
have nuchal constriction; 26 genera.

Genus Tintinnopsis Stein, 1867


Reference: Kofoid and Campbell, 1929

Codonellidae with lorica usually bowl-shaped, never with a narrow oral aperture; distinct aboral
end closed or rarely with an irregular (broken?) aperture; sometimes with a posterior horn but more
usually not; wall thin and homogeneous, or with a fine primary structure, with freely agglomerated
foreign matter as well as particles of detritus, scattered abundantly on outer surface; spiral structure
rare.
This is a very complex genus of many species practically all of which are either neritic, or fresh-
water inhabitants. The characterization of the species of this genus is very difficult and as yet, the
systematics of the group are not satisfactory, largely because of the great irregularities of the loricae
due to agglomeration, which obscures the underlying structure apparent in most other genera.
Differs from Codonella in less regular pattern of the secondary structure of the wall and in the lack of
a nuchal groove or an inner nuchal shelf.

Tintinnopsis acuminata Daday, 1887


(Fig. 5 a-d)

Description: Lorica tubular, oral rim ragged, aboral region in lower 1/4 conical (70 degrees), aboral
end blunt. Wall without spiral structure, with spare agglomeration.

Reported Measurements: Total length 54-78 µm. Oral diameter 18-21 µm. Approximate ratio L/oral
diameter 2.9-3.5.

Measurements of Specimens from Kuwait Waters: Length 60-86 µm; oral diameter 23-30 µm;
approximate ratio L/oral diameter 2.3-3.2. According to Daday’s description, oral region has no oral
funnel; in the samples some T. acuminata have good visible oral expansion (Fig. 5 b).

Distribution: Tintinnopsis acuminata was registered in relatively low abundance in the northwestern
Arabian Gulf.

20 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

Fig. 5 a-d. Variations of lorica shape of Tintinnopsis acuminata. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 21


Tintinnopsis ampla Hada, 1937
(Fig. 6 a-d)

Description: Lorica comparatively large, generally bullet-shaped; ratio of lorica length to oral
diameter is 1.8-2.6; oral margin entire; bowl cylindrical or very slightly tapering (up to 7 degrees);
aboral region tapering abruptly (77-100 degrees), rarely having a short aboral horn; aboral end blunt
or acute; wall thickened, composed of rather coarse agglomerated materials, spiral structure visible,
when well-developed sides of the bowl becoming uneven.

Comments: This species is remarkably variable in form, represented by several different forms. The
general contour of most individuals is gradually tapering in the bowl, straight on sides of the lorica,
and lacking an aboral horn, but some are rugged in appearance on account of the well-developed
spiral structure and of the presence of a stout aboral horn. In specimens without an aboral horn, the
aboral end is typically blunt or acute and occasionally rounded. There are a few intermediate forms.
This species differs from T. beroidea Stein in being of larger size, from T. japonica Hada in the conical
aboral region and the coarser agglomeration of the wall, and from T. elongata Daday in the presence
of the spiral structure and the shape of the aboral region. Elongated conical individuals of the
species closely resemble T. elongata in general outline, but are distinguishable from the latter by the
presence of a spiral structure and by the abruptly narrowed aboral region.

Reported Measurements: Length 157 (130-192) µm; oral diameter 73 (70-75) µm.

Measurements of Specimens from Kuwait Waters: Length 201 (160-260) µm; oral diameter 116
(110-120) µm; approximate ratio L/oral diameter 1.5-2.6.

Distribution: In the cool season T. ampla is the most abundant species in the waters surrounding the
Bubiyan Island. Abundance of this species reaches to 94.5 X 103 organisms/m3, with 34% percentage
composition of total abundance of tintinnids in February 2004 (Skryabin and Al-Yamani, 2007a).
Previously, T. ampla was registered only in Japanese waters of the Pacific Ocean. Probably, this
species was introduced with ballast waters of oil tankers into Kuwait waters and became successfully
established.

22 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

Fig. 6 a-d. Variations of lorica shape of Tintinnopsis ampla. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 23


Tintinnopsis angusta Meunier, 1910
(Fig. 7 a-c)

Description: Lorica short, rather fusiform but oral diameter more than half the maximum diameter
of bowl.

Reported Measurements: Length 70 µm; oral diameter 20 µm; approximate ratio L/oral diameter
3.5.

Measurements of Specimens from Kuwait Waters: Length 68-75 µm; oral diameter 32-45 µm;
greatest transdiameter 39-54 µm; approximate ratio L/oral diameter 1.7-2.1.

Distribution: Tintinnopsis angusta was registered in relatively low abundance in the northwestern
waters of the Arabian Gulf.

a b

Fig. 7 a-c. Variations of lorica shape of Tintinnopsis angusta. Scale – 50 µm.

24 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Tintinnopsis baltica Brandt, 1986
(Fig. 8 a-d)

Description: Lorica campanulate, 1.6-2.5 greatest transdiameters in length; aboral margin flaring,
roughened with agglomerated materials, its oral diameter 1.6-2.5 of the greatest transdiameter;
bowl more or less inflated in the posterior 0.4-0.6 of the lorica; aboral region conical (50-80 degrees),
occasionally tapering to an inconspicuous stout aboral horn; aboral end usually bluntly pointed;
aboral opening usually lying laterally (Fig. 8 c), often comparatively large; wall rather coarsely
agglomerated, sometimes with a few slight spiral turns in the suboral region.

Comments: This species differs from T. brevicollis in dimension, and the smaller bowl opening
aborally, and from T. akkeshiensis in the presence of the posterior dilation.

Reported Measurements: Length 50-80 µm; oral diameter 30-43 µm; greatest transdiameter 30-37
µm; approximate ratio L/oral diameter 1.7-1.9

Measurements of Specimens from Kuwait Waters: Length 57-73 µm; oral diameter 30-46 µm;
greatest transdiameter 36-42 µm; approximate ratio L/oral diameter 1.6-1.9.

Distribution: Tintinnopsis baltica was registered in low abundance in the northwestern waters of the
Arabian Gulf.

a b

c d

Fig. 8 a-d. Variations of lorica shape of Tintinnopsis baltica. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 25


Tintinnopsis beroidea Stein, 1867
(Fig. 9 a-c).

Description: Lorica bullet-shaped, usually cylindrical in the anterior 0.6-0.7 of the total length,
aborally conical (75-85 degrees), oral rim ragged; aboral end acute or bluntly pointed; wall rather
coarse, 0.03-0.04 oral diameters in thickness, without spiral structure.

Comments: The species differs from T. acuminata Daday in stouter proportions and in the subacute
aboral end, from T. parvula Jorgensen in the absence of an aboral expansion below the anterior
subcylindrical region, and from T. strigosa Meunier in the aboral end.

Reported Measurements: Total length 34-110 µm. Oral diameter 18-40 µm. Approximate ratio of
L/oral diameter 1.5-3.0.

Measurements of Specimens from Kuwait Waters: Length 60-80 µm; oral diameter 27-36 µm;
greatest transdiameter 31-35 µm; approximate ratio L/oral diameter 1.9-2.2.

Distribution: Tintinnopsis beroidea was registered in relatively low abundance in the northwestern
waters of the Arabian Gulf.

a
b

Fig. 9 a-c. Variations of lorica shape of Tintinnopsis beroidea. Scale – 50 µm.

26 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Tintinnopsis compressa Daday, 1887
(Fig. 10 a, b).

Description: Lorica small, rounded, with slight nuchal constriction and oral expansion. Wall thin,
sparse agglomerated.

Reported Measurements: Total length 47-90 µm. Oral diameter 37-75 µm. Approximate ratio L/oral
diameter 1.0-1.3.

Distribution: Tintinnopsis compressa was registered in low abundance in the northwestern waters
of the Arabian Gulf.

a b

Fig. 10 a-b. Variations of lorica shape of Tintinnopsis compressa. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 27


Tintinnopsis dadayi Kofoid, 1905
(Fig. 11 a-h).

Description: Lorica elongated, consisting of a low oral flare, a cylindrical collar, and a globose bowl,
approximately 0.7 greatest transdiameters of the bowl in length; oral rim formed irregularly, about
greatest transdiameters of the bowl in oral diameter; aboral end hemispherical or bluntly pointed;
wall coarsely agglomerated.

Reported Measurements: Total length 65-81 µm. Oral diameter 43-49 µm. Minimum diameter of
collar 29-31 µm; greatest transdiameter of bowl 37-42 µm. Approximate ratio L/oral diameter 1.5-
1.7.

Measurements of Specimens from Kuwait Waters: Total length 70-80 µm; oral diameter 47-51 µm;
minimum diameter of collar 36-44 µm; greatest transdiameter of bowl 45-50 µm; length of collar 40-
56 µm; length of bowl 30-34 µm; approximate ratio L/oral diameter 1.5-1.6. In Kuwait waters loricas
often have secondary oral rims (Figs. 10 d, h).

Distribution: Tintinnopsis dadayi is common in the northwestern waters of the Arabian Gulf.

a b

28 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


c d

e f

g h

Fig. 11 a-h. Variations of lorica shape of Tintinnopsis dadayi. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 29


Tintinnopsis directa Hada, 1932
(Fig. 12 a-f ).

Description: Lorica tall campanulate, oral diameter is 1.6-2.2 µm in length; oral rim irregular, flaring
(60-92 degrees); suboral region somewhat tapering, conical (5-10 degrees), laid up with about 6
spiral turns, narrowest at the basal portion of the subcylindrical part, its smallest transdiameter is
0.68-0.82 of the oral diameter; posterior region subspherical, with a rounded aboral end, 0.80-0.95
oral diameter in transdiameter; wall rather coarse in the posterior part, is about 0.035 diameters in
thickness at the thickest portion of the aboral region.

Comments: The species differs from T. everta Kofoid and Campbell and T. cyathus Daday in having
a distinct globose posterior region, from T. everta Kofoid and Campbell in having a distinct aboral
enlargement, from T. pallida Brandt in the presence of the more differentiated aboral part, and from
T. turgida Kofoid and Campbell in the possession of a flare of the oral rim.

Reported Measurements: Total length 72-100 µm; oral diameter 35-48 µm; greatest transdiameter
of the bowl 32-40 µm. Approximate ratio L/oral diameter 2.1-2.7. Some T. directa in Kuwait waters
have slightly developed bowl (Fig. 11 c).

Distribution: Tintinnopsis directa is common and abundant in the northwestern waters of the
Arabian Gulf and in the central region of the Gulf, mainly along the Iranian coast.

30 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

f
e

Fig. 12 a-f. Variations of lorica shape of Tintinnopsis directa. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 31


Tintinnopsis failakkaensis Skryabin and Al-Yamani, 2006
Reference: Skryabin and Al-Yamani, 2006
(Fig. 13 a-g).

Description: Lorica comparatively large, pot-shaped; oral diameter is 0.67-0.77 of total length; bowl
convex, aboral end usually bluntly pointed (Figs. 13 a, b, c), some specimens have rounded (Figs.
13 d, e) or sharply pointed (Figs. 13 f ) aboral end. Oral flare relatively wide, campanulate, the angle
with the longitudinal axis of lorica is 35-45 degrees, in upper half is 0-10 degrees. Lower part of flare
usually has from five to nine impressions; that is why on drawing and photographs, usually one side
of lower part of oral flare looks less smooth than the other side. Bowl often has one or two deep
spiral dents (45-55 degrees). The length of flare is about 1/4 - 1/6 total length of lorica.

Comments: This tintinnid has lorica of Codonella-like shape. However, representatives of the genus
Codonella have wall of lorica with Cittarocylis structure (Kofoid and Campbell, 1929), with few or
no agglomerated particles, but often with coccoliths (Marshall, 1969). The new species has walls
without any Cittarocylis structure, densely covered with coarse Tintinnopsis-like agglomeration. Few
coccoliths are present only on collar, never on bowl. The reliable difference between these two
closely related genera is the structure of the wall, not the shape of lorica because some species of
Tintinnopsis (i.e. T. orientalis, T. schotti, T. mortensenii, etc.) also have well-differentiated neck and bowl
and a conspicuous nuchal constriction.
This species differs from T. shotti, T. dadayi, and T. everta in bigger size, campanulate oral flare and
comparatively more wide, convex and shortened bowl and from T. orientalis in longer bowl, more
campanulate collar and bigger size.

Measurements of Specimens from Kuwait Waters: Total length of lorica 137 µm (126-149 µm);
diameter of oral flare 99 µm (93-103 µm); diameter of nuchal constriction 76 µm (67-82 µm); greatest
transdiameter of bowl 85 µm (77-93 µm); approximate ratio of L/oral diameter is 1.3-1.5.
Scatter plots of lorica length against oral diameter of T. failakkaensis and T. orientalis based on
microscopic measurements of 50 specimens for each species are displayed in Fig. 13 g. The rest of
the 24 species of this genus found in Kuwait waters fell out of the scale of the above figure.

Distribution: In cool season T. failakkaensis was usually observed in waters surrounding Failakka
Island and outside the entrance of Kuwait Bay. Maximum density (up to 3.0 X 103 organisms/m3) was
recorded in December 2003 near the southern coast of Failakka Island.

a b

Fig. 13. Tintinnopsis failakkaensis. Scale – 50 µm.


a-f. variations of lorica shape; g. scatter plots of lorica length against oral diameter and linear regression between
length and maximum width of lorica (with 99% confidence interval) of T. failakkaensis and T. orientalis based on
measurements of 50 specimens of each species.

32 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


c d

e f

110
T. failakkaensis T. orientatis

105

100
Diameter of oral flare (µm)

95

90

85

80

90 100 110 120 130 140 150 160

Length of lorica (µm)


g

Fig. 13. Cont’d

K u wait I n s titute f or Scienti f ic R e s earc h 33


Tintinnopsis gracilis Kofoid and Campbell, 1929
(Fig. 14 a-d).

Description: Lorica finger-shaped, oral margin usually comparatively smooth; bowl tubular,
sometimes slightly swollen in the posterior one-third of the bowl; aboral region

convex conical (40-75 degrees) with a blunt distal end; wall coarsely agglomerated without a spiral
structure.

Comments: The species differs from T. karajacensis Brandt in the conical aboral region instead of the
round.

Reported Measurements: Length 105-125 µm; oral diameter 28-34 µm; approximate ratio L/oral
diameter 3.3-4.0.

Measurements of Specimens from Kuwait Waters: Length 109-130 µm; oral diameter 36-45 µm;
greatest transdiameter 42-46 µm; approximate ratio L/oral diameter 2.8-3.2.

Distribution: Tintinnopsis gracilis is one of the most common and abundant species in the
northwestern Arabian Gulf and in the central region of the Gulf, mainly along the Iranian coast.

34 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

Fig. 14 a-d. Variations of lorica shape of Tintinnopsis gracilis. d. conjugation of two T. gracilis organisms.
Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 35


Tintinnopsis karajacensis Brandt, 1896
(Fig. 15).

Description: Lorica capsular, bowl cylindrical in the margin part, oral rim ragged; aboral end rounded
or disfigured as the result of irregularly agglomerated particles; wall coarse, having several slight
spiral turns in the anterior half.

Comments; This species differs from T. directa Hada in the absence of the oral flare and the
posterior inflation, from T. cochleata (Brandt) in less extensive spiral organization and in roughened
agglomeration, from T. lobiancoi Daday in the shorter lorica, and from T. rotundata Jorgensen in
more slender proportions and in the shape of the aboral end.

Reported Measurements: Length 60-172 µm; oral diameter 30-64 µm; approximate ratio L/oral
diameter 2.0-2.7

Measurements of Specimens from Kuwait Waters: Length 107-115 µm; oral diameter 35-38 µm;
approximate ratio L/oral diameter 3.0.

Distribution: Tintinnopsis karajacensis was registered in relatively low abundance in the northwestern
Arabian Gulf.

Fig. 15. Lorica shape of Tintinnopsis karajacensis. Scale – 50 µm.

36 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Tintinnopsis lobiancoi Daday, 1887
(Fig. 16 a-d).

Description: Lorica elongate, tubular, usually straight, oral rim ragged; aboral end rounded or
shaped somewhat irregularly; wall agglomerated roughly, but comparatively thin, 0.04 of the oral
diameter in thickness, without a spiral structure.

Comments: Differs from T. karajacensis Brandt in having longer lorica and more slender proportions
and from T. cochleata (Brandt) in the lack of the spiral structure.

Reported Measurements: Length 93-409 µm; oral diameter 34-62 µm; approximate ratio L/oral
diameter 2.0-7.0.

Measurements of Specimens from Kuwait Waters: Length 96-115 µm; oral diameter 28-31 µm;
greatest transdiameter 34-37 µm; approximate ratio L/oral diameter 3.5.

Distribution: Tintinnopsis lobiancoi was registered in relatively low abundance in the northwestern
Arabian Gulf.

a b

c d

Fig. 16 a-d. Variations of lorica shape of Tintinnopsis lobiancoi. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 37


Tintinnopsis lohmanni Laackmann, 1906
(Fig. 17 a-b).

Description: Lorica vase-like with a cylindrical collar; collar usually short, 0.16-0.40 of the total length
in length; bowl expanding to its greatest transdiameter of 1.06-1.24 oral diameters; aboral region
rounded or convex conical; wall coarsely agglomerated, a few spiral turns appearing in the collar.

Comments: Differs from T. subacuta Jörgensen in proportion of the collar to the bowl and from T.
tubulosa Levander in posterior inflation and the presence of the spiral structure.

Reported Measurements: Length 72 (58-110) µm; oral diameter 48 (35-52) µm; length of the collar
20 (10-50) µm; greatest transdiameter of the bowl 55 (40-62) µm; approximate ratio L/oral diameter
1.3-2.2.

Distribution: Tintinnopsis lohmanni is abundant in the cool season in the waters surrounding
Bubiyan Island.

a b

Fig. 17 a-b. Variations of lorica shape of Tintinnopsis lohmanni. Stained with Rose Bengal. Scale – 50 µm.

38 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Tintinnopsis mortensenii Schmidt, 1901
(Fig. 18 a-d).

Description: Lorica bell-shaped with slightly concave collar, very wide, horizontal oral funnel and
rounded aboral end.

Reported Measurements: Length 90-110 µm; oral diameter 87-100 µm; approximate ratio L/oral
diameter 1.0-1.1.

Measurements of Specimens from Kuwait Waters: Length 50-70 µm; oral diameter 52-92 µm;
greatest transdiameter of bowl 33-45 µm; minimum transdiameter of collar 26-35 µm; approximate
ratio L/oral diameter 0.9-1.5.

Distribution: Tintinnopsis mortensenii is common in the northwestern waters of the Arabian Gulf
and in the central region of the Gulf, mainly along the Iranian coast.

a b

Fig. 18 a-d. Variations of lorica shape of Tintinnopsis mortensenii. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 39


Tintinnopsis nana Lohmann, 1908
(Fig. 19 a-d).

Description: Lorica very small, cylinder with slightly pointed aboral end. Oral rim often ragged.
Wall thin, with poor agglomeration.

Comments: This species differs from T. beroidea Stein in being of smaller size and in the shape
of the aboral region, and differs from T. minuta only in shape of aboral end and ratio of length to
oral diameter, both of which are rather variable characters.

Reported Measurements: Length 30-47 µm; oral diameter 12-19 µm; approximate ratio L/oral
diameter 1.7-3.5.

Measurements of Specimens from Kuwait Waters: Length 29-39 µm; oral diameter 21-29 µm;
approximate ratio L/oral diameter 1.3-1.6. In Kuwait’s samples T. nana had wider lorica, often with
expanded oral end (Fig. 19 c).

Distribution: Tintinnopsis nana is common in the northwestern Arabian Gulf, in the central
region of the Gulf along the Iranian coast, and near the Strait of Hormuz.

a b

c d

Fig. 19 a-d. Variations of lorica shape of Tintinnopsis nana. Scale – 20 µm.

40 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Tintinnopsis orientalis Kofoid and Campbell, 1929
(Fig. 20 a-d).

Description: Lorica with distinct collar; collar 1/4-1/3 length of the bowl in length, convex
outwardly; bowl acorn-shaped, widest near or shortly below its middle; aboral end baggy, not
flattened, with or without a faintly emergent point.

Comments: Differs from T. dadayi in a convex, instead of concave bowl; widely flaring collar and
less baggy, less flattened aboral end.

Reported Measurements: Length 90-110 µm; oral diameter 87-100 µm; approximate ratio L/
oral diameter 1.0-1.1.

Measurements of Specimens from Kuwait Waters: Length 106 µm (98-113 µm); oral diameter
85 µm (82-87 µm); greatest transdiameter of lorica 79 µm (75-82 µm); diameter of nuchal
constriction 70 µm (67-72 µm); approximate ratio L/oral diameter 1.1-1.3. In Kuwait waters
majority of loricas have irregularly formed aboral opening usually set laterally (Figs. 20 a, b, c).

Distribution: Tintinnopsis orientalis is relatively common in the northwestern waters of the


Arabian Gulf during the warm season.

a b

c d

Fig. 20 a-d. Variations of lorica shape of Tintinnopsis orientalis. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 41


Tintinnopsis parva Merkle, 1909
(Fig. 21 a-c).

Description: Lorica small, widest in middle, narrowing slightly to mouth, more sharply to pointed
aboral end.

Reported Measurements: Length 30-49 µm; oral diameter 20-23 µm; greatest transdiameter of the
bowl 15-31 µm; approximate ratio L/oral diameter 1.2-2.5.

Measurements of Specimens from Kuwait Waters: Length 50-65 µm; oral diameter 25-30 µm;
greatest transdiameter 43-46 µm; approximate ratio L/oral diameter 1.2-1.5.

Distribution: Tintinnopsis parva is relatively common in the northwestern waters of the Arabian
Gulf.

a b

Fig. 21 a-c. Variations of lorica shape of Tintinnopsis parva. Scale – 50 µm.

42 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Tintinnopsis parvula Jörgensen, 1912
(Fig. 22 a-d).

Description: Lorica small, slightly expanded below a cylindrical anterior region, pointed aborally.
Comments: Lorica more pointed aborally than T. nucula and more expanded than T. beroidea.

Reported Measurements: Length 51-70 µm; oral diameter 20-37 µm; approximate ratio L/oral
diameter 1.8-3.4.

Measurements of Specimens from Kuwait Waters: Length 62-85 µm; oral diameter 27-35 µm;
greatest transdiameter 32-42 µm; approximate ratio L/oral diameter 2.0-2.5.

Distribution: Tintinnopsis parvula is common and abundant in the northwestern waters of the
Arabian Gulf and in the waters surrounding Bubiyan Island.

a b

c d

Fig. 22 a-d. Variations of lorica shape of Tintinnopsis parvula. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 43


Tintinnopsis radix Imhof, 1886
(Fig. 23 a-f ).

Description: Lorica elongate, slender, tubular, oral rim generally entire (smooth and round) or
sometimes irregular; bowl long, cylindrical; aboral region tapering gradually into an aboral horn,
inverted conical (41-21 degrees); aboral horn, usually more or less curved, with an irregularly formed
aboral opening typically set laterally as gouged, leaving its tip or cutting off it; wall thin and fragile,
0.03 of the oral diameter in thickness, with a variable spiral structure.

Comments: The species differs from T. kofoidi Hada in the fragile construction of the lorica, in less
contraction at the aboral region, and in the shape of the lateral opening in the aboral horn, and from
T. cylindrica in open aboral end and spiral structure of lorica.

Reported Measurements: Total length 140-524 µm. Oral diameter 32-75 µm. Approximate ratio L/
oral diameter 6.0-9.5.

Measurements of Specimens from Kuwait Waters: Length 195-500 µm; oral diameter 45-95 µm;
approximate ratio L/oral diameter 4.3-7.0.

Distribution: Tintinnopsis radix is common and abundant at all the studied locations in the Arabian
Gulf, from Bubiyan Island in the north to the Strait of Hormuz in the south.

44 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

e f

Fig. 23 a-f. Variations of lorica shape of Tintinnopsis radix. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 45


Tintinnopsis rotundata Jörgensen, 1899
(Fig. 24).

Description: Lorica cylindrical with hemispherical aboral end; oral margin usually ragged; wall
comparatively thin, thickly encrusted with particles, spiral structure invisible.

Comments: This species differs from the typical form of T. karajacensis Brandt in the possession of
the thin wall without spiral structure.

Reported Measurements: Total length 40-90 µm. Oral diameter 24-47 µm. Approximate ratio L/oral
diameter 1.5-2.1.

Measurements of Specimens from Kuwait Waters: Length 80 µm; oral diameter 26 µm;
approximate ratio L/oral diameter 3.1.

Distribution: Tintinnopsis rotundata was registered in low abundance in the northwestern waters
of the Arabian Gulf.

Fig. 24. Lorica shape of Tintinnopsis rotundata. Scale – 50 µm.

46 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Tintinnopsis sacculus Brandt, 1896
(Fig. 25 a-d).

Description: Lorica short, cylindrical, with rounded aboral end; particles smaller and less numerous
than on T. rotundata.

Reported Measurements: Length 60-105 µm; oral diameter 44-58 µm; approximate ratio L/oral
diameter 1.5-2.0.

Measurements of Specimens from Kuwait Waters: Length 50-65 µm; oral diameter 41-47 µm;
approximate ratio L/oral diameter 1.2 -1.4.

Distribution: Tintinnopsis sacculus was registered in low abundance in the northwestern waters of
the Arabian Gulf.

a b

c d

Fig. 25 a-d. Variations of lorica shape of Tintinnopsis sacculus. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 47


Tintinnopsis schotti Brandt, 1906
(Fig. 26 a-d).

Description: Lorica bell-shaped, constricted in the suboral 1/4-1/5 of the total length; oral rim
irregular; collar widely flaring to form an inverted truncated low cone of 45-75 degrees with convex
sides, its shortest basal diameter 0.68-0.78 of an oral diameter; bowl cup-shaped, usually broadest
a little below its middle, 0.75-0.85 of an oral diameter in greatest diameter; aboral region broadly
convex conical (96-120 degrees) to a blunt distal end; wall coarsely agglomerated, thickened to
make an inward projection at the nuchal constriction, no spiral structure.

Comments: The species differs from T. orientalis Kofoid and Campbell in having a more flaring low
collar and a more distinct nuchal constriction and from T. loricata Brandt in possessing a stouter
bowl.

Reported Measurements: Total length 96-120 µm. Oral diameter 82-85 µm. Nuchal diameter 58-65
µm. Greatest diameter of bowl 62-70 µm. Approximate ratio L/oral diameter 1.1-1.4.

Measurements of Specimens from Kuwait Waters: Length 56-69 µm. Oral diameter 48-60 µm.
Nuchal diameter 36-37 µm. Greatest diameter of bowl 39-41 µm. Approximate ratio L/oral diameter
1.1-1.2. In Kuwait waters, loricas of T. schotti rarely have secondary oral rim (Fig. 26 d).

Distribution: Tintinnopsis schotti is common in the northwestern waters of the Arabian Gulf.

48 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

Fig. 26 a-d. Variations of lorica shape of Tintinnopsis schotti. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 49


Tintinnopsis tocantinensis Kofoid and Campbell, 1929
(Fig. 27 a-f ).

Description: Lorica elongated, cylindrical anteriorly, expanding posteriorly, tapering distally into a
stout aboral horn; dilated part not spiraled, 1.2 of the oral diameter in transdiameter; aboral horn
conical (35 degrees), obliquely or irregularly open at the tip (Fig. 27 f, g); wall thick and coarse.

Comments: Differs from Tintinnopsis aperta Brandt in the absence of the spiral structure at the
enlarged region and in having the stout aboral horn.

Reported Measurements: Total length 103 µm. Oral diameter 22 µm. Approximate ratio L/oral
diameter 4.7.

Measurements of Specimens from Kuwait Waters: Length 82-143 µm; oral diameter 17-22 µm;
greatest transdiameter 27-33 µm; approximate ratio L/oral diameter 4.7-6.5.

Distribution: Tintinnopsis tocantinensis is common in the northwestern waters of the Arabian Gulf.

50 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

d
c

e
f

Fig. 27 a-d. Variations of lorica shape of Tintinnopsis tocantinensis; e. aboral end of lorica; f. conjugation of two
specimens of T. tocantinensis. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 51


Tintinnopsis turbo Meunier, 1919
(Fig. 28).

Description: Lorica small, rounded with a bluntly pointed aboral end. Widest in the middle,
narrowing to the oral end, which may have short cylindrical section.

Reported Measurements: Total length 54-63 µm. Oral diameter 37-38 µm. Approximate ratio L/oral
diameter 1.6-1.7.

Measurements of Specimens from Kuwait Waters: Length 55-68 µm; oral diameter 36-38 µm;
greatest transdiameter 43-49 µm; approximate ratio L/oral diameter 1.5-1.8.

Distribution: Tintinnopsis turbo was registered in low abundance in the northwestern Arabian Gulf.

Fig. 28. Tintinnopsis turbo. Scale – 50 µm.

52 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Tintinnopsis undella Meunier, 1910
(Fig. 29).

Description: Lorica vase-shaped, slightly expanded in lower 2/3, widening slightly to mouth.

Reported Measurements: Length 75-94 µm; oral diameter 20-38 µm; greatest transdiameter of the
bowl 42 µm; approximate ratio L/oral diameter 2.6.

Measurements of Specimens from Kuwait Waters: Length 78-85 µm; oral diameter 35-36 µm;
greatest transdiameter 40-43 µm; approximate ratio L/oral diameter 2.2-2.3.
Distribution: Tintinnopsis undella was registered in low abundance in the northwestern Arabian
Gulf.

Fig. 29. Tintinnopsis undella. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 53


Tintinnopsis urnula Meunier, 1910
(Fig. 30 a-d).

Description: Lorica campanulate; oral rim ragged; bowl constricted at the suboral 1/3 of the total
length, its narrowest transdiameter is 0.8 oral diameter, sides concave in the anterior 2/3 of the
lorica; aboral region an inverted cone of 78 degrees; aboral end subacute; wall more or less coarse,
with very slight spiral organization in the suboral part.

Comments: Differs from T. beroidea Stein and T. lata Meunier in the presence of the suboral
constriction, and from T. compressa Daday in the widened oral aperture and in the tapering aboral
end.

Reported Measurements: Length 54-60 µm; oral diameter 37-40 µm; approximate ratio L/oral
diameter 1.5.

Measurements of Specimens from Kuwait Waters: Length 60-75 µm; oral diameter 50-55 µm;
greatest transdiameter 40-45 µm; minimum diameter of collar 36-39 µm; approximate ratio L/oral
diameter 1.2-1.4.

Distribution: Tintinnopsis urnula is common in the northwestern waters of the Arabian Gulf.

54 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

Fig. 30 a-d. Variations of lorica shape of Tintinnopsis urnula. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 55


FAMILY: CODONELLOPSIDAE Kofoid and Campbell, 1929

Lorica top-shaped, thick walled; agglomerate of mineral particles; 12 genera.

Genus Codonellopsis Jörgensen, 1924


Reference: Kofoid and Campbell, 1929

Codonellopsidae with lorica divided into an annular collar and a bowl; collar set off distinctly in
structure from bowl, often narrower than the bowl, hyaline with spiral structure, or at least with one
or two coiled bands, and even these sometimes obscured; bowl oval to spheroidal, with or without
hollow antapical horn closed off from the bowl by a basal membrane; wall of bowl with primary,
secondary and tertiary structure, sometimes with “fenestrae”; agglomerating material often on wall,
as Tintinnopsis structure.
Differs from Stenosemella in the higher collar and from Laackmaniella in the closed instead of open
aboral end.

Codonellopsis lusitanica Jörgensen, 1924


Reference: Marshall, 1969
(Fig. 31 a-d).

Description: Collar usually a little longer than bowl with a few spiral turns at oral end. Bowl narrow,
bluntly pointed aborally. Particles scattered thickly on bowl, thinly on collar.

Reported Measurements: Length 94 µm; oral diameter 23 µm; greatest transdiameter of the bowl
34 µm; approximate ratio L/oral diameter 4.1.

Distribution: Codonellopsis lusitanica is common and abundant in the northwestern waters of the
Arabian Gulf and in the waters surrounding Bubiyan Island. Maximum density (up to 3.4 million
organisms/m3, 89.2% of total tintinnid’s abundance) was recorded in February 2005 near the
southern coast of Bubiyan Island.

56 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

Fig. 31 a-d. Variations of lorica shape of Codonellopsis lusitanica. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 57


Codonellopsis morchella (Cleve, 1900)
Reference: Hada, 1937
(Fig. 32 a-d).

Description: Lorica 2.2-3.2 oral diameters in length; collar with a somewhat flaring rim, slightly
bulging near its middle, usually lower than the bowl, its height 0.22-0.34 of the total length, composed
of 4-11 spiral turns with a few elliptical fenestrae; bowl generally ovate, 1.5-1.7 oral diameters in
greatest diameter; aboral region convex conical; aboral end usually round or rarely bluntly pointed;
wall of the bowl thick, coarsely agglomerated.

Comments: The species differs from C. americana in its shorter lorica and ovate bowl instead of an
ellipsoid one and from C. schabi in its low bowl.

Reported Measurements: Length 65-95 µm; oral diameter 29-32 µm; length of the collar 14-32 µm;
greatest diameter of the bowl 43-65 µm. Approximate ratio L/oral diameter 2.2-3.2.

Distribution: Codonellopsis morchella is common in the northwestern and central regions of the
Arabian Gulf and in the Strait of Hormuz, with high abundance in the central region of the Gulf.

a b

c d

Fig. 32 a-d. Variations of lorica shape of Codonellopsis morchella. Stained with Rose Bengal. Scale – 50 µm.

58 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Codonellopsis ostenfeldi (Schmidt, 1901)
Reference: Kofoid and Campbell, 1929
(Fig. 33 a-d).

Description: Lorica flask-shaped, oral diameter is 0.37-0.33 of total length; oral margin entire
(smooth and round), slightly flaring; collar subcylindrical, 0.40-0.43 of the total length in height, with
2-4 spiral turns in its anterior end and 5-6 (in Kuwait waters often to 12-15) annular rows of elliptical
fenestrae lying transversely on the lower part (Fig. 33 d); bowl globose, its greatest transdiameter 1.6
oral diameters; aboral end broadly rounded; wall agglomerated with coarse foreign particles.

Comments: The species differs from the other species of Codonellopsis in having spiral rows of
numerous fenestrae on the collar.

Reported Measurements: Length 95-112 µm; oral diameter 35-38 µm; length of the collar 38-50
µm; greatest transdiameter of the bowl 55-62 µ.

Measurements of Specimens from Kuwait Waters: Length of this species reaches to 227 µm (bowl
up to 87 µm, collar up to 140 µm) in Kuwait waters.

Distribution: Codonellopsis ostenfeldi is common and abundant in the northwestern and central
regions of the Arabian Gulf and in the Strait of Hormuz.

b
a

c
d

Fig. 33 a-c. Variations of lorica shape of Codonellopsis ostenfeldi; d. structure of collar. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 59


FAMILY: DICTYOCYSTIDAE Haeckel, 1873
Reference: Lynn and Small, 2000.

Lorica collar with one or two rows of quadrangular fenestrae; coccoliths or mineral particles as
foreign material; all marine; two genera.

Genus Dictyocysta Ehrenberg, 1854


Reference: Kofoid and Campbell, 1929

Dictyocystidae with lorica with a tall cylindrical collar with beams surrounding large windows in
one to several rows; bowl rounded, inflated; wall with coarse and fine reticulations and often with
included coccoliths.

Dictyocysta duplex Brandt, 1906


References: Kofoid and Campbell, 1929; Marshall, 1969
(Fig. 34 a-b).

Description: Lorica with oral margin undulating; collar 0.3 – 0.5 total length, with a single row of
7-9 tall rectangular windows whose length is 1.2 of their width; beams sometimes bowed outwards.
Bowl subglobular, or rarely wider (0.8 length of bowl) than long, set off by distinct shoulder from
the bowl; aboral region subhemispherical. Wall of bowl with large, subuniform, coarse, rounded and
overlapping meshes and with smaller ones intermingled, formed by included coccoliths. Fenestrae
sometimes present, variable in pattern (Fig. 34 b).

Comments: The species differs from the other species of Dictyocysta in the extensive coccolith
inclusions and duplex nature of the wall structure.

Reported Measurements: Length 53-75 µm; oral diameter 32-42 µm; approximate ratio L/oral
diameter 1.6-2.3.

Distribution: Dictyocysta duplex is abundant in the waters of the Strait of Hormuz.

60 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a

Fig 34 a-b. Variations of lorica shape of Dictyocysta duplex. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 61


Genus Luminella Kofoid and Campbell, 1939

Genus Luminella takes the intermediate position between two genera – Stenosemella and
Dictyocysta and has some differences from each of them. Stenosemella has bowl densely covered
with agglomerated particles and short hyaline cylindrical collar without windows. Luminella has
very close shape of lorica, but differs in presence of windows in the collar. Dictyocysta has collar
with beams surrounding windows, but never has bowl densely covered with agglomerates. Thus,
Luminella probably is the evolutionary stage of development of comparatively primitive lorica of
Stenosemella into more progressive lorica of Dictyocysta.
Kofoid and Campbell (Marshall, 1969) described the genus Luminella on the basis of a single species
(L. parvicollis), which they reported as a new species. Since a second species was encountered in
Kuwait’s samples, it was deemed necessary to modify the description of the genus.

Modified Description of the Genus Luminella Kofoid and Campbell, 1939

Collar very short, hyaline, with beams, or with small semicircular windows at base. Bowl globose or
cup-shaped, bluntly pointed or hemispherical aborally. Surface covered with particles.

Luminella kuwaitensis Skryabin and Al-Yamani, 2007


References: Al-Yamani and Skryabin, 2006; Skryabin and Al-Yamani, 2007.
(Fig. 35 a-f ).

Description: Collar short, approximately 1/10 of total length, hyaline consists of 8-10 thin connected
arcs enclosing big squarish fenestrations (Figs. 35 a-f ). Bowl cup-shaped (Figs. 35 a, c, d, f ), heart-
shaped (Fig. 35 b), or globose (Fig. 35 e), bluntly pointed or hemispherical aborally, usually forms
pronounced shoulder below collar. Surface densely covered with big agglomerated particles. Bowl
is without any visible concentric structures.

Comments: L. kuwaitensis differs from L. parvicollis in its smaller size of lorica, collar formed with 8-10
arcs, not cylindrical with 12 small windows and absence of concentric striae on the bowl.

Measurements of Specimens from Kuwait Waters: Total length 52.1 µm (47.5-58.8 µm). Length
of bowl 47.0 µm (42.5-50.0 µm). Maximum width of bowl 47.2 µm (42.5-52.5 µm). Diameter of oral
aperture 26.8 µm (22.0-32.5 µm). Length of collar 5.1 µm (5.0-7.5 µm). Width of separate arch 8.6 µm
(8.0-10.0 µm).

Distribution: L. kuwaitensis is one of the most abundant species in Kuwait waters and the north-
western part of the Arabian Gulf. It was found in this area at all stations in all seasons, but with
lower numbers in winter. Abundance of this species reaches 6.7 million organisms/m3, with
70% percentage composition of total number of tintinnids recorded in October, 2003 (average
percentage composition for studied area during August-October, 2003 was 56% of total tintinnid
abundance). In Kuwait Bay in December 2007, when water temperature fell below 18°C, mass
conjugation of L. kuwaitensis was observed (Fig. 35 e). This species changed the asexual type of
propagation into sexual reproduction, and produced resting stages, and in January 2008 almost
completely disappeared at the studied area until the beginning of the warm season.

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a b

c d

e f

Fig. 35 a-e. Variations of lorica shape of Luminella kuwaitensis; f. conjugation of L. kuwaitensis. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 63


FAMILY: METACYLIDIDAE Kofoid and Campbell, 1929
Reference: Lynn and Small, 2000.

Lorica tubular or globet-shaped, with rings or spiral band on all or part, wall 3-layered, usually with
distinct alveoli; 10 genera.

Genus Coxliella Brandt, 1908


Reference: Kofoid and Campbell, 1929

Tintinnida with lorica generally tall bowl- or vase-like; oral rim never regularly denticulate; without
a well differentiated collar; wall double, usually with two laminae, with coarse secondary structure;
lorica formed throughout or in greater part by a single spiral band with superposed turns of varying
heights.
Some authors consider the genus Coxliella invalid, since the genus Favella when deprived of lorica
builds new (secondary) lorica with coxliella-like shape (Laval-Peuto, 1977).

Coxliella annulata (Daday, 1886)


Reference: Marshall, 1969
(Fig. 36)

Description: Lorica tubular but very slightly wider toward aboral end, bluntly pointed aborally.
Spiral turns slightly overlapping. Wall structure indistinct.

Comments: C. annulata usually was registered together with F. campanula and could be a
representative of F. campanula when forming a secondary lorica.

Reported Measurements: Total length 269-332 µm; oral diameter 100-128 µm; approximate ratio
L/oral diameter 2.7-3.0.

Measurements of Specimens from Kuwait Waters: Total length 225 µm; oral diameter 112 µm;
approximate ratio L/oral diameter 2.0.

Distribution: Coxliella annulata was observed in low abundance in the northwestern Arabian Gulf.

64 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Fig. 36. Lorica shape of Coxliella annulata. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 65


Genus Helicostomella Jörgensen, 1924
Reference: Kofoid and Campbell, 1929

Tintinnida with elongate, narrow lorica; suboral region cylindrical, formed at least in part by a
spirally wound band of 3-60 turns; aboral region conical; aboral horn present; wall thin, with a
fine uniform primary structure between the two lamellae.

Helicostomella longa Kofoid and Campbell, 1929


Reference: Kofoid and Campbell, 1929
(Fig. 37 a-d)

Description: Lorica very short, bullet-shaped, 2.2-4.4 oral diameters in length; oral rim entire; 4-11
subequal suboral turns, not everted orally; bowl expanding slightly, widest 0.5-1.0 oral diameter
below the spiral, convex conical (40-70 degrees) aborally; aboral horn scarcely differentiated,
conical (35 degrees).

Comments: H. longa differs from all other species in having shorter, stouter lorica.

Reported Measurements: Total length 50-80 µm.

Measurements of Specimens from Kuwait Waters: Total length 50-60 µm; maximum width
16-17 µm; length of collar 17-19 µm (0.3-0.4 total length), approximate ratio L/oral diameter 3.1-
3.5.

Distribution: Helicostomella longa was registered in very low abundance in the northwestern
waters of the Arabian Gulf.

66 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

Fig. 37 a-d. Variations of lorica shape of Helicostomella longa. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 67


Genus Metacylis Jörgensen, 1924
Reference: Kofoid and Campbell, 1929

Metacylididae with lorica usually short and wide, oval, in some species elongate capsular; usually
with a wide mouth and a short wide collar with a few, low spiral turns or annuli; bowl wide; aboral
end rounded or pointed, flattened, or with a spinule; wall bilaminate, simple with indistinct structure,
or with simple prismatic structure, or hyaline.

Metacylis jorgensenii (Cleve, 1902)


(Fig. 38 a-d)

Description: Lorica short, ovoid, with slightly or sharply pointed aboral end. Collar short, with 2-5
spiral turns, cylindrical or slightly flaring, narrower than bowl.

Reported Measurements: Total length 50-61 µm; oral diameter 44-50 µm; approximate ratio L/oral
diameter 1.3-1.9.

Measurements of Specimens from Kuwait Waters: Total length 61-67 µm; oral diameter 55-56 µm;
greatest transdiameter 63-64 µm; approximate ratio L/oral diameter 1.1-1.2.
Distribution: Metacylis jorgensenii usually is relatively rare in Kuwait waters and is observed mainly
at the end of the warm season, during August-September.

68 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

Fig. 38 a-d. Variations of lorica shape of Metacylis jorgensenii. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 69


Metacylis lucasensis Kofoid and Campbell, 1929
(Fig. 39 a-b)

Description: Small species with a cylindrical, capsule-like lorica, collar with four spiral turns in
the anterior 0.3 of total length; oral margin entire; aboral end hemispherical, without point; wall
very thin. Differs from all other species in the cylindrical shape of the lorica.

Reported Measurements: Total length (L) 47 µm; oral diameter 27 µm; approximate ratio L/oral
diameter 1.75.

Measurements of Specimens from Kuwait Waters: Total length 51-56 µm; oral diameter 35-36
µm; approximate ratio L/oral diameter 1.5-1.6.

Distribution: Metacylis lucasensis was registered in relatively low abundance in the northwestern
waters of the Arabian Gulf.

70 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a

Fig. 39 a, b. Variations of lorica shape of Metacylis lucasensis. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 71


Metacylis pithos Skryabin and Al-Yamani, 2006
Reference: Skryabin and Al-Yamani, 2006
(Fig. 40 a-g)

Description: Lorica small, hyaline, pot-shaped, shape of aboral end varies from bluntly to sharply
pointed. Collar cylindrical, relatively short (0.14-0.23 of total length), with 1-6 spiral turns. Diameter
of collar varies from 80 to 97% of maximum diameter of bowl.

Comments: M. pithos differs from M. pontica in cylindrical collar, from M. jorgensenii and M.
mereschkovskii in proportions, from M. rossica in smaller size and in M. corbula in pointed aboral
end.

Measurements of Specimens from Kuwait Waters: Total length 38-57 µm; maximum transdiameter
32-35 µm; oral diameter 28-31 µm, length of collar 8-10 µm, approximate ratio L/oral diameter 1.5-
2.0.
Scatter plots of lorica length against greatest transdiameter of lorica of M. pithos and M. tropica
based on microscopic measurement of 100 specimens of M. pithos and 50 specimens of M. tropica
are displayed in Fig. 40 g.

Distribution: Metacylis phitos is common at all studied area of the Arabian Gulf, from Bubiyan Island
to the Strait of Hormuz with high abundance in the northwestern region.

a b

Fig. 40. Metacylis pithos. Scale – 50 µm.


a-f. variations of lorica shape; g. scatter plots of lorica length against greatest transdiameter
diameter, and linear regression between length and maximum width of lorica (with 99%
confidence interval) of M. pithos and M. tropica based on measurements of 100 specimens of
M. pithos and 50 specimens of M. tropica.

72 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


c

e f

60
M. tropica
M. pithos
105
Greatest trandiameter of lorica (µm)

50

45

40

35

30

25
35 40 45 50 55 60 65

Length of lorica (µm) g


Fig. 40. Cont’d

K u wait I n s titute f or Scienti f ic R e s earc h 73


Metacylis tropica Duran, 1957
(Fig. 41 a-c)

Description: Lorica small, global, length is approximately equal to maximum width. Collar with
3-5 spiral turns, 0.14-0.30 of the total length of lorica.

Reported Measurements: Total length 43 µm; oral diameter 33-40 µm; maximum diameter 41-
44 µm.

Distribution: Metacylis tropica is a common species in the northwestern waters of the Arabian
Gulf. In February 2009, very high abundance of M. tropica (up to 1.0-1.1 million organisms/m3;
91.9% - 96.1% of total tintinnid’s abundance and 91.4% - 96.8% of total biomass) was registered
along the southern coast of Kuwait Bay.

a b

Fig. 41 a-c. Variations of lorica shape of Metacylis tropica. Scale – 50 µm.

74 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Metacylis sp.
(Fig. 42 a-b)

Description: Lorica small, hyaline, cup-shaped, with rounded aboral end. Collar wide, relatively
short, with 4-6 spiral turns.

Comments: These tintinnids are close to M. pithos in size and shape, but differs in widely opened
oral aperture. It could be a form of M. pithos.

Measurements of Specimens from Kuwait Waters: Length 39-41 µm; oral diameter 35-39 µm;
greatest transdiameter 36-40 µm; approximate ratio L/oral diameter 1.0-1.1.

Distribution: Metacylis sp. is relatively rare in the northwestern waters of the Arabian Gulf.

Distribution: Metacylis sp. was registered in low abundance in the northwestern waters of the
Arabian Gulf.

b
a

Fig. 42 a-b. Variations of lorica shape of Metacylis sp. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 75


FAMILY: ASCAMPBELLIELLIDAE Corliss, 1960
Reference: Marshall, 1969; Lynn and Small, 2000.

Very small forms, cup or tube shaped with rounded or pointed aboral end. Circumoral region
differentiated into inner collar and outer collar or flaring lip with gutter or slope between. Wall
trilaminate with intermediate layer hyaline or weakly alveolar. Five genera.

Genus Ascampbelliella Corliss, 1960


Reference: Marshall, 1969

Ascampbelliellidae with very small tubular or cup-shaped bowls. Circumoral region always with
two rims: an inner collar and flaring lip. Outer rim sometimes repeated. No teeth.

Ascampbelliella obscura (Brandt, 1906)


(Fig. 43 a-c)

Description: Lorica cone-shaped, narrowing more abruptly in aboral ¼ to a sharp point. Inner
collar erect, outer slightly flaring, with gutter between. Well marked alveolar structure in wall.

Reported Measurements: Total length 70-90 µm; oral diameter 42-53 µm; approximate ratio L/
oral diameter 1.6-1.7.

Distribution: Ascampbelliella obscura was observed at all studied area of the Arabian Gulf with
highest abundance in the western part of the Strait of Hormuz.

76 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a

Fig. 43 a-c. Variations of lorica shape and structure of lorica of Ascampbelliella obscura. Stained
with Rose Bengal. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 77


FAMILY: PETALOTRICHIDAE Kofoid and Campbell, 1929
Reference: Marshall, 1969; Lynn and Small, 2000.

Lorica bowl-shaped with round or pointed aboral end and collar in two parts, suboral cone and
flaring lip. Nuchal constriction usually present. Small, suboral and subnuchal fenestrae on lip and
bowl. Two genera.

Genus Petalotricha Kent, 1881


Reference: Kofoid and Campbell, 1929

Petalotrichidae with bowl-shaped or conical lorica, oral shelf spreading; low oral ridge, flaring
conical collar; slight or deep nuchal constriction, prominent internal nuchal ledge; globular,
sack-like, or subconical bowl; aboral end hemispherical, flattened, minutely pointed or bluntly
pointed; wall with inner and outer laminae enclosing 1-3 layers of minute, faint or thick-walled,
rounded, uneven prisms; suboral fenestrae in one row with horizontal axis, subnuchal fenestrae
circular or elliptical in a more or less restricted zone, with long axis oblique or vertical.

Petalotricha ampulla (Fol, 1881)


(Fig. 44 a-b)

Description: Bowl globose or ovoid, with rounded or pointed aboral end. Suboral cone nearly
cylindrical, lip flaring. Edge of lip often irregularly denticulate. Ring of small oval fenestrae at top
of cone and a double row on shoulder of bowl.

Reported Measurements: Total length 116-165 µm; oral diameter 100-135 µm; approximate
ratio L/oral diameter 1.1-1.6.

Distribution: Petalotricha ampulla was registered in central and eastern regions of the Arabian
Gulf.

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a

Fig. 44 a-b. Variations of lorica shape of Petalotricha ampulla. Stained with Rose Bengal. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 79


FAMILY: RHABDONELLIDAE Kofoid and Campbell, 1929
Reference: Lynn and Small, 2000.

Lorica acorn- or chalice-shaped with longitudinal, low ridges that may be simple, branch or
anostomose; four genera.

Genus Protorhabdonella Jorgensen, 1924


Reference: Kofoid and Campbell, 1929

Rhabdonellidae with lorica short and stout; oral aperture never with teeth; always simple; bowl
more or less conical; with few or many vertical or subvertical ribs; rarely with an aboral horn; wall
single, usually hyaline or with primary prisms only.
Differs from Rhabdonella in the absence of the oral gutter and from Rhabdonellopsis in the
absence of pedicel and knob.

Protorhabdonella curta (Cleve, 1901).


(Fig. 45 a-b)

Description: Lorica with more or less conical bowl, slightly dilated above middle. Aboral end
pointed, closed. Oral rim simple, with only one slight thickening of wall below. Ribs 18-24, vertical
or slightly spiral; no branching. No fenestrae. Wall thin hyaline.

Comments: P. curta differs from P. simplex in having more numerous ribs and smaller size and
from P. mira and P. striatura in the absence of pedicel.

Reported Measurements: Total length 39-52 µm; oral diameter 22-29 µm, approximate ratio L/
oral diameter 1.4-1.8.

Distribution: Protorhabdonella curta was registered in the central and eastern regions of the
Arabian Gulf.

a b

Fig. 45 a-b. Variations of lorica shape of Protorhabdonella curta. Stained with Rose Bengal. Scale – 50 µm.

80 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Protorhabdonella simplex (Cleve, 1900).
(Fig. 46 a-c)

Description: Lorica short, convex-conical, with low collar and pointed aboral end. Ribs 6-10,
vertical, no branching. Wall thick just below collar.

Comments: P. simplex differs from P. curta in having less ribs and bigger size, and from P. mira and
P. striatura in the absence of pedicel.

Reported Measurements: Total length 47-98 µm; oral diameter 26-39 µm, approximate ratio L/
oral diameter 1.5-2.2.

Distribution: Protorhabdonella simplex was registered in the central and eastern regions of the
Arabian Gulf.

a b

Fig. 46 a-c. Variations of lorica shape of Protorhabdonella simplex. Stained with Rose Bengal. Scale – 50 µm

K u wait I n s titute f or Scienti f ic R e s earc h 81


Genus Rhabdonella Brandt, 1906
Reference: Kofoid and Campbell, 1929

Rhabdonellidae usually with elongated chalice-shaped lorica; oral rim never with teeth,
but usually with a gutter between the inner and outer laminae; pedicel slender, more or less
protracted, never with apophyses; ribs more or less vertical, simple or branched and sometimes
with anastomoses, with few to many or no fenestrae between the ribs, usually in the suboral
region; laminae well developed, especially toward the mouth, generally with one layer of primary
prisms and usually a coarse Cyttarocylis structure.
Differs from Protorhabdonella and Rhabdonellopsis in the presence of pedicel and knob.

Rhabdonella conica Kofoid and Campbell, 1929


(Fig. 47 a-d)

Description: Lorica very long, chalice-shaped, 5.6-7.9 oral diameters in length; oral rim flaring
abruptly, no higher than lip; bowl tapering conical (upper 10-28 degrees, lower 30-50 degrees), it
is about half the length of the total length of the whole lorica; pedicel very long, subcylindrical,
tapering distally; aboral end open; ribs 32-48, with sinistral deflection, almost vertical on top half,
slightly spiral to near aboral end; fenestrae distinct, widely distributed.

Comments: This species differs from R. spiralis and R. striata in having more conical form of the
bowl and in more gradual transition between bowl and pedicel.

Reported Measurements: Total length 290-470 µm. Oral diameter 37-102 µm. Approximate
ratio L/oral diameter 4.8-7.9.

Distribution: Rhabdonella conica is very rare in the northwestern waters of the Arabian Gulf. This
species was registered in Summer 2003 near Kubbar Island and in Spring and Summer 2009 in
coastal waters of Kuwait and in Kuwait Bay.

82 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

Fig. 47. Variations of lorica shape of Rhabdonella conica. a-d. variations of lorica shape; e – structure of lorica.
Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 83


Rhabdonella striata (Biedermann, 1893)
(Fig. 48 a-c)

Description: Lorica cylindrical-conical with slender pedicel more than 0.3 total length. Oral rim
hardly emergent, lip flaring. Wall with very distinct laminae, thick in pedicel. Ribs 24-54, vertical
or slightly spiral. Fenestrae small, numerous.

Comments: This species is shorter and stouter than R. spiralis, longer than R. brandti.

Reported Measurements: Total length 200-410 µm. Oral diameter 40-57 µm. Approximate ratio
L/oral diameter 3.3-4.6.

Distribution: Rhabdonella striata is very rare in the northwestern waters of the Arabian Gulf. This
species was registered only in Summer 2003 near Kubbar Island.

a b

Fig. 48. Variations of lorica shape of Rhabdonella striata. a-b. variations of lorica shape; c. structure of lorica.
Scale – 50 µm.

84 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Rhabdonella sp.
(Fig. 49 a-b)

Description: Lorica almost cylindrical above and conical (7°-9°) below. Oral rim flaring, relatively
wide, no higher than lip. Pedicel very short, 0.2 total length, with small roundish knob or swelling
on the tip. Ribs almost vertical on oral half, curving slightly right to near aboral end. Fenestrae
numerous.

Comments: The species differs from the other species of Rhabdonella in wide flaring oral rim and
short pedicel with swelling on tip.

Measurements of Specimens from Arabian Gulf: Total length 155-185 µm. Oral diameter 45-
50 µm. Approximate ratio L/oral diameter 3.6-3.7.

Distribution: Rhabdonella sp. was registered in the eastern region of the Arabian Gulf.

Fig. 49. Lorica shape of Rhabdonella sp. a. shape of lorica; b. structure of lorica. Stained with Rose Bengal.
Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 85


FAMILY: XYSTONELLIDAE Kofoid and Campbell, 1929
Reference: Lynn and Small, 2000.

Lorica chalice-shaped; irregular and conspicuous meshwork between wall layers with
inconspicuous, thin wall material separating polygons; six genera.

Genus Parundella Jörgensen, 1924


Reference: Kofoid and Campbell, 1929

Xystonellidae with elongated lorica with an upper cylinder and a lower cone prolongated usually
as a simple caudal lance; wall with a simple primary structure, or hyaline, its laminae separated
but approximate, except at the aboral end where they fuse.
Differs from Xystonella in the absence of the channeled oral rim and from Xystonellopsis in having
the thin, not flaring oral rim.

Parundella lohmanni Jörgensen, 1924


(Fig. 50 a-b)

Description: Lorica more or less cylindrical, but dilated just below mouth, contracted in middle
of bowl, and expanding again slightly before thick pedicel. Wall thick, lamellae well separated
except where they fuse in tip of pedicel.

Reported Measurements: Total length 151-183 µm. Oral diameter 38-43 µm. Approximate ratio
L/oral diameter 3.9-4.3.

Distribution: Parundella lohmanni is common in the central and eastern parts of the Arabian
Gulf with highest abundance recorded in the eastern regions.

a b

Fig. 50 a-b. Variations of lorica shape of Parundella lohmanni. Stained with Rose Bengal. Scale – 50 µm.

86 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Genus Xystonella Brandt, 1907
Reference: Kofoid and Campbell, 1929

Xystonellidae with lorica subconical, very long and narrow; oral margin thickened and channeled
between the two lamellae, the outer lamella more or less flaring, and an inner usually forming
an erect collar; bowl elongate, vase-like or conical; prolonged aborally in a spine or in pedicel
with knob or apophyses; with or without terminal lance; wall with a single middle prismatic layer.
Differs from Xystonellopsis and Parundella in having the oral rim channeled.

Xystonella treforti (Daday, 1887)


(Fig. 51 a-c)

Description: Lorica long, cylindrical-to vase-shaped, with pedicel ending in skirt, which has 6-8
folds and points. Slender lance, varying in width. Inner collar low, lip slightly flaring and may be
denticulate.

Reported Measurements: Total length 275-500 µm. Oral diameter 40-96 µm. Approximate ratio
L/oral diameter 4.4-9.1.

Distribution: Xystonella treforti was registered in low abundance in the central part of the
Arabian Gulf.

a b

Fig. 51. Lorica shape of Xystonella treforti. a. shape of


lorica; b. shape of aboral end and skirt; c. structure of
bowl. Stained with Rose Bengal. Scale – 50 µm
c

K u wait I n s titute f or Scienti f ic R e s earc h 87


Genus Xystonellopsis Jörgensen, 1924
Reference: Kofoid and Campbell, 1929

Xystonellidae with lorica elongated and usually narrow, chalice-shaped; oral rim thinning to
sharp edge without a gutter; bowl vase-shaped or conical; pedicel very elongated, often with
skirt; terminal lance usually present; wall with three well developed lamellae, the middle one of
numerous small prismatic alveoli.
Differs from Xystonella in the absence of the channel in the oral rim and from Parundella in having
an abruptly flared oral margin.

Xystonellopsis gaussi (Laackmann, 1909)


(Fig. 52)

Description: Lorica relatively short and stout, no suboral ring; bowl 0.75 total length in length,
inverted subconical (3°) anteriorly; aboral cone contracting more rapidly (20°-22°), 1.0 oral
diameter in length; pedicel 0.6-1.0 oral diameter in length; terminating in a squarely truncate
skirt without expansion; with 6-8 dexiotropic lists on pedicel and lower part of cone; lance
slender, conical (15°), 0.4-0.5 oral diameter in length.

Comments: Differs from X. dahli in having a skirt.

Reported Measurements: Total length 371-470 µm. Oral diameter 60-70 µm. Approximate ratio
L/oral diameter 5.6-6.6.

Distribution: Xystonellopsis gaussi was registered in low abundance in the central part of the
Arabian Gulf, along the Iranian coast.

Fig. 52. Lorica shape of Xystonellopsis gaussi. Stained with Rose Bengal. Scale – 50 µm.

88 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Genus Favella Jörgensen, 1924
Reference: Kofoid and Campbell, 1929

Xystonellidae with lorica generally campanulate or subconical; oral rim entire or with small skirt, or with
denticles, with or without a suboral constriction and ridge; never with a collar separated with a band from the
bowl proper, but sometimes with one or more annuli; bowl campanulate to conical, contracting aborally; aboral
horn present, generally thick-walled; wall bilamellate, usually with coarse, intermediate prismatic secondary
alveoli and a very fine, primary structure, newer with regular polygonal structure. Only marine species.
Differs from Protocymatocylis and Cymatocylis in the absence of striae; from Poroecus in better developed
more distinctly prismatic wall and more suboral development; and from Parafavella in much less regular, less
distinct, and smaller prisms in the wall.

Favella adriatica (Imhof, 1886) Jörgensen emended


(Fig. 53 a-c)

Description: Lorica cylindrical, bell-shaped, with pedicel 1/4-2/3 total length. Wings sometimes visible.

Reported Measurements: Total length 170-336 µm. Oral diameter 112-128 µm. Approximate ratio L/oral
diameter 1.6-2.6.

Distribution: Favella adriatica was registered in low abundance in the northwestern Arabian Gulf.

a b

Fig. 53 a-c. Variations of lorica shape of Favella


adriatica. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 89


Favella campanula (Schmidt) Jörgensen, 1924
(Fig. 54 a-c)

Description: Lorica campanulate, oral rim more or less irregular; bowl with a raised band just
below the suboral ring, cylindrical in its upper 0.65; aboral region convex conical (80°) to a stout
horn; aboral horn having a few vertical striae on the surface, tapering (20°) to a bluntly pointed
tip, its length 0.15 of the total length; wall almost hyaline with a hardly visible reticulation,
apparently separated in the suboral inflated part and scarcely in the following 0.15 of the total
length.

Comments: The species differs from F. azorica (Cleve) in the presence of an aboral horn.

Reported Measurements: Length 143 µm; oral diameter 68 µm. Approximate ratio L/oral
diameter 2.1.

Distribution: Favella campanula is common and abundant in the northwestern Arabian Gulf
during the warm period.

a b

Fig. 54 a-c. Variations of lorica shape of Favella campanula. Scale – 50 µm.

90 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Favella ehrenbergi (Claparede and Lachmann) Jörgensen emended
(Fig. 55 a-c)

Description: Lorica long, cylindrical; bowl sometimes slightly expanded below middle, rounded
below and joined by wings to a short blunt, pedicel. Spiral turns sometimes present suborally.
Wall thick.

Reported Measurements: Total length 145-400 µm. Oral diameter 54-124 µm. Approximate
ratio L/oral diameter 2.4-4.2.

Distribution: Favella ehrenbergi was registered in low abundance in the northwestern waters of
the Arabian Gulf.

a
b

Fig. 55 a-c. Variations of lorica shape of Favella ehrenbergi. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 91


Favella panamensis Kofoid and Campbell, 1929
(Fig. 56 a-f )

Description: Lorica cylindrical, stout, oral rim entire, with 1, rarely 2-4 rings, slightly lipped below
the ring; bowl without nuchal constriction, subcylindrical for 0.60-0.75 its length; aboral region
contracting abruptly; aboral horn 0.3-0.8 oral diameter in length, conical (20 degrees), with
oblique wings, tip pointed.

Comments: Differs from F. ehrenbergi in having a more cylindrical bowl and fuller aboral region.

Reported Measurements: Total length 136-232 µm. Oral diameter 37-38 µm. Approximate ratio
L/oral diameter 2.0-2.9.

Distribution: Favella panamensis is common and abundant in the northwestern waters of the
Arabian Gulf during the cool period.

92 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

e
f

Fig. 56 a-f. Variations of lorica shape of Favella panamensis. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 93


FAMILY: UNDELLIDAE Kofoid and Campbell, 1929
Reference: Lynn and Small, 2000.

Lorica goblet-shaped; clear wall; inner and outer layers of wall, conspicuously thick; eight genera.

Genus Undella Daday, 1887


Reference: Kofoid and Campbell, 1929

Undellidae with lorica with the suboral region tapering in section to a thin oral margin; without
suboral ledge or inner collar; bowl cylindrical or nearly so anteriorly, never greatly or abruptly
expanded aborally, without rings; aboral end rounded, angled, pointed or flattened; wall
bilaminate, without reticulations and without prismatic structure.
Differs from Undellopsis in the absence of the suboral ledge, from Cricundella and Amplectella in
the absence of rings, from Proplectella in the absence of the inner collar, and from Amplectellopsis
in the absence of the bowl.

Undella dilatata Kofoid and Campbell, 1929


(Fig. 57 a-d)

Description: Lorica subcylindrical, expanding posteriorly to slightly inflated, faintly angular


aboral region; suboral region not expanded beyond the diameter of the bowl, narrowing in a
sigmoid curve on the outer face for 0.1-0.2 oral diameter below the rounded oral rim; cylinder
with constant diameter for 0.6-0.7 total length, then bulging to 1.1-1.4 oral diameters at about 0.7-
0.8 total length, the upper slope of the bulge varying in length and convexity but not distinctly
angular; aboral end subconical (125°-150°), broadly to acutely pointed; wall subuniformin
thickness (0.1 oral diameter) in the cylinder, thickest in lower part of the bulge (0.15) and thinnest
(0.05) in the aboral point.

Comments: The species differs from the other species of U. hyalina group un the dilated aboral
region and from U. dohrnii in the lack of anterior contraction of the bowl.

Reported Measurements: Length 130-160 µm; oral diameter 48-52 µm. Approximate ratio L/
oral diameter 2.5-3.3.

Distribution: Undella dilatata was registered in low abundance in the central part of the Arabian
Gulf.

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a b

c d

Fig. 57 a-d. Variations of lorica shape of Undella dilatata. Stained with Rose Bengal. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 95


FAMILY: TINTINNIDAE Claparéde and Lachmann, 1858
Reference: Lynn and Small, 2000.

Lorica long, oral end flared; wall 2-layered, homogeneous, clear: inner and outer layers of wall
not thick; 23 genera.

Genus Amphorellopsis Kofoid and Campbell, 1929


Reference: Hada, 1937

Lorica usually elongate, consisting of a low funnel-shaped collar and a subcylindrical or fusiform
bowl with pointed end; oral margin usually entire; aboral region tapering to an aboral end, from
which a number of vertical fins or ridges sometimes extend to the collar; wall single-layered
except an anterior flaring part of the lorica (Hada, 1937).

Amphorellopsis acuta (Schmidt) Kofoid and Campbell, 1929


(Fig. 58 a-d)

Description: Lorica fusiform, oral aperture circular; collar low-funnel-shaped (70-85 degrees),
its nuchal smallest diameter 0.54-0.60 of an oral diameter; bowl circular in cross section below
the collar, then gradually becoming triangular, posteriorly with three ridges in the aboral 0.6 of
the total length; aboral end acute; wall composed of separated laminae in the anterior 0.2 of the
lorica.

Comments: The species differs from the other species of Amphorellopsis in having three ridges.

Reported Measurements: Length 85-108 µm; oral diameter 31-45 µm. Approximate ratio L/oral
diameter 2.3-3.9.

Measurements of Specimens from Kuwait Waters: Length 92-102 µm; oral diameter 36-49 µm;
approximate ratio L/oral diameter 2.1-2.7.

Distribution: Amphorellopsis acuta is common at all studied areas of the Arabian Gulf.

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a
b

c d

Fig. 58 a-d. Variations of lorica shape of Amphorellopsis acuta. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 97


Genus Amphorides Strand, 1926
Reference: Hada, 1937

Lorica tubular or vase-shaped, usually with flaring collar. Oral rim entire (or toothed), aboral end
closed, often truncated. Bowl with 3, 4, or 8 longitudinal rides or fins. Wall hyaline, structureless.

Amphorides amphora (Claparede and Lachmann, 1858)


(Fig. 59 a-e)

Description: Lorica with flaring collar; oral aperture circular; usually with flaring collar. Greatest
width in lower part of bowl, with truncated aboral end. Three low vertical fins. Wall thickened at
neck.

Reported Measurements: Length 100-220 µm; oral diameter 55 µm, approximate ratio L/oral
diameter 2.5-4.0.

Measurements of Specimens from Kuwait Waters: Length 100-244 µm; oral diameter 45-65
µm; approximate ratio L/oral diameter 3.4-4.5.

Distribution: Amphorides amphora is common at all studied areas of the Arabian Gulf.

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a b

c d

Fig. 59. Amphorides amphora. a-d. variations of lorica shape; e. structure of aboral end. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 99


Amphorides quadrilineata Jorgensen, 1924
(Fig. 60 a-c)

Description: Lorica tall urn-shaped; collar low funnel-shaped, its basal diameter 0.68-0.78 oral
diameters; bowl circular in cross section in the anterior part, but gradually triangular in the
posterior on account of development of three conspicuous fins extended through the entire
bowl; aboral end truncate, more or less concave in its center. Variable in form, greatest width
above or below middle of bowl.

Comments: Because the lower part of the lorica is triangular, frontal and side views of a specimen
are different. The species differs from A. brandti in having a stouter form and from A. infundibulum
in the structure of the collar and fins.

Reported Measurements: Length 92-182 µm; oral diameter 38-66 µm; approximate ratio L/oral
diameter 2.4-3.6.

Distribution: Amphorides quadrilineata was common at all the studied area of the Arabian Gulf.

a b

Fig. 60 a-c. Variations of lorica shape of Amphorides


quadrilineata. Stained with Rose Bengal. Scale – 50 µm.

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Genus Dadayiella Kofoid and Campbell, 1929
Reference: Kofoid and Campbell, 1929

Tintinnida with an elongated goblet-shaped lorica, 2.6-5.8 oral diameters in length; oral rim
crenulated or entire; collar with 9-18 oblong facets; bowl campanulate, usually contracting
posteriorly to an elongated conical or cylindrical pedicel with or without an aboral knob; and
with or without a pointed aboral tip.

Dadayiella cuspis Kofoid and Campbell, 1929


(Fig. 61 a-b)

Description: Lorica small, elongated; collar flaring 12 degrees, oral facets 11-13 on both collar
and bowl extending to the pedicel; bowl subcylindrical conical posteriorly; pedicel short, 0.1-0.2
total length.

Reported Measurements: Total length (L) 67-93 µm; oral diameter 22-31 µm. Approximate ratio
L/oral diameter 2.9-3.1.

Measurements of Specimens from Kuwait Waters: The oral facetes of lorica of all specimens
from Kuwait waters sticking above the oral rim and form long spines, about 0.1-0.2 length of
lorica.
Total length 92-102 µm; oral diameter 30-31 µm; approximate ratio L/oral diameter 3.1-3.3.

Distribution: Dadayiella cuspis was relatively common at all the studied areas of the Arabian
Gulf.

a b

Fig. 61 a-b. Variations of lorica shape of Dadayiella cuspis. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 10 1


Dadayiella ganymedes (Entz, 1884)
(Fig. 62 a-b)

Description: Lorica elongate, almost tubular or with a slightly flaring oral region, tapering from
about ½ total length to slender pedicel, about1/4 total length, pointed or obtuse at aboral tip.
Pedicel often has 9-12 low longitudinal fins throughout its length. Oral region facetted with 9-18
ribs, sometimes sticking up above oral rim. Nine secondary ribs sometimes appear between nine
stronger primary ribs.

Comments: Y. Hada (1937) also included in this species D. acutiformes, D. bulbosa and D.
jorgenseni.

Reported Measurements: Total length 75-120 µm; oral diameter 25-30 µm. Approximate ratio
L/oral diameter 3.0-4.2.

Distribution: Dadayiella ganymedes was registered in central and eastern regions of the Arabian
Gulf.

a b

Fig. 62 a-b. Variations of lorica shape of Dadayiella ganymedes. Stained with Rose Bengal. Scale – 50 µm.

10 2 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Genus Eutintinnus Kofoid and Campbell, 1939
Reference: Kofoid and Campbell, 1939

Tintinnidae with lorica in the form of a truncated cone or cylinder open at both ends; wall homogeneous,
hyaline, rarely externally wrinkled, never with spiral structure.
Differs from all other genera of Tintinnidae in the lack of all structural differentiations in the oral and aboral ends,
except those produced by eversion, by the thickening of the margins into a brim, and by the presence of teeth.

Eutintinnus apertus (Kofoid and Campbell, 1929)


(Fig. 63 a-c)

Description: Lorica chalice-shaped; oral region slightly if at all flaring; bowl subconical (15°-20°) anteriorly,
slightly concave laterally; aboral region contracting abruptly to 0.3-0.6 oral diameter; aboral end truncated,
open. Often with laterally attached pelagic diatoms, especially Chaetoceros.

Comments: This species differs from E. angustatus in wider aboral region.


E. apertus from the Arabian Gulf often has an asymmetrical lorica. Aboral aperture of the Gulf specimens is
shifted sideways relatively to the central axis of the lorica (Fig. 63 c).

Reported Measurements: Length 85-112 µm; oral diameter 28-39 µm, approximate ratio L/oral diameter 2.2-3.2.

Distribution: Eutintinnus apertus was registered in central and eastern regions of the Arabian Gulf.

a b

Fig. 63 a-c. Variations of lorica shape of Eutintinnus


apertus. Stained with Rose Bengal. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 10 3


Eutintinnus conicus Skryabin and Al-Yamani, 2006
Reference: Skryabin and Al-Yamani, 2006.
(Fig. 64 a-b)

Description: Lorica has a shape of a short truncated cone, narrowing about 4 – 5 degrees. Oral
end flaring to a slightly emergent brim. Aboral end strait. Aboral diameter little less than 1/2 oral.

Comments: The species differs from E. lusus-undae and E. tenue in shorter lorica and more conical
shape. In Kuwait waters, difference between these three species is visible (Fig. 65 e).

Measurements of Specimens from Kuwait Waters: Length 166-178 µm; oral diameter 42-48
µm; aboral diameter 21-23 µm; approximate ratio oral/aboral diameter 2.0-2.1; approximate ratio
L/oral diameter 3.7.

Distribution: Eutintinnus conicus was registered in low abundance in the northwestern waters
of the Arabian Gulf.

a b

Fig. 64 a, b. Eutintinnus conicus. Scale – 50 µm.

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Zone I
Zone II

Zone III

Zone IV

The different zones sampled in the Arabian Gulf during 1999-2010


- Zone I: Waters around Bubiyan Island
- Zone II: Kuwait Bay
- Zone III: Northwestern region of the Arabian Gulf
- Zone IV: Central and Eastern Arabian Gulf

K u wait I n s titute f or Scienti f ic R e s earc h 10 5


Eutintinnus contractus Skryabin and Al-Yamani, 2006
Reference: Skryabin and Al-Yamani, 2006
(Fig. 65 a-e)

Description: Lorica very long and slender. Oral flare, 5-6 degrees, takes approximately 20% of
total length; the greatest part of lorica, about 60% is nearly tubular; 1-2 degrees, aboral narrowing
(6 degrees) takes 15% of total length, after narrowest point lorica sharply (26-27 degrees), flared
to aboral end (Fig. 65 e).

Comments: The species differs from another species of Eutintinnus in the shape of aboral part
of lorica.

Measurements of Specimens from Kuwait Waters: Length 433-461 µm oral diameter 60-65
µm; aboral diameter 24-27 µm; aboral narrowing 15-20 µm; approximate ratio L/oral diameter
7.0-7.4.
Scatter plots of lorica length against oral diameter of E. conicus, E. lusus-undae and E. contractus
based on microscope measurements of 50 specimens of each species are displayed in Fig. 65 e.

Distribution: Eutintinnus contractus was registered in low abundance in the northwestern waters
of the Arabian Gulf.

a b

Fig. 65. Eutintinnus contractus. Scale – 50 µm.


a-c. variations of lorica shape; d. aboral end; e. scatter plots of lorica length against oral diameter and linear
regression between length and maximum width of lorica (with 99% confidence interval) of E. conicus, E. contractus
and E. lusus-undae based on measurements of 50 specimens of each species.

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c d

70
E. Iusus-undae
E. contractus
65 E. conicus

60
Oral diameter (µm)

56

50

45

40

35
100 150 200 250 300 350 400 450 500

Length of lorica (µm) e


Fig. 65. Cont’d

K u wait I n s titute f or Scienti f ic R e s earc h 10 7


Eutintinnus fraknoi (Daday, 1887)
(Fig. 66)

Description: Lorica large, tapering, gently flaring both orally and aborally. Aboral diameter about
2/3 that of oral. Oral brim present. Occasionally with polygonal prisms and included particles in
wall. Sometimes with attached Chaetoceros.

Reported Measurements: Length 278-475 µm; oral diameter 40-80 µm, approximate ratio L/
oral diameter 4.5-6.5.

Measurements of Specimens from Kuwait Waters: Length 260-301 µm; oral diameter 46-54
µm; aboral diameter 25-30 µm; approximate ratio L/oral diameter 5.5-6.0.

Distribution: Eutintinnus fraknoi was registered in low abundance in the northwestern waters of
the Arabian Gulf.

Fig. 66. Eutintinnus fraknoi. Scale – 50 µm.

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Eutintinnus lusus-undae (Entz, 1885) (from Marshall, 1969)
(Fig. 67 a-d)

Description: Lorica almost cylindrical with slightly flaring oral end and low brim. No flare or brim
aborally. Aboral diameter little more than 1/2 oral. Wall sometimes with prisms, sometimes with
attached Chaetoceros.

Reported Measurements: Total length 169-290 µm. Oral diameter 38-54 µm. Approximate ratio
L/oral diameter 3.2-4.9.

Measurements of Specimens from Kuwait Waters: Length 255-265 µm; oral diameter 50-55
µm; aboral diameter 25-26 µm; approximate ratio L/oral diameter 5.0.

Distribution: Eutintinnus lusus-undae is relatively common at all studied areas of the Arabian
Gulf.

a b

c d

Fig. 67 a-d. Variations of lorica shape of Eutintinnus lusus-undae. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 10 9


Eutintinnus macilentus (Jörgensen, 1924)
(Fig. 68 a-c)

Description: Lorica a short truncated cone (6-9 degrees); oral margin with a bream; shaft flaring
(18-29 degrees) anteriorly, contracting as a cone (3-5 degrees) toward the aboral end to a least
diameter, 0.5 oral diameter, at a level of 1.0 oral diameter from the aboral end; flaring aborally
(30 degrees).

Reported Measurements: Length 136-290 µm; oral diameter 37-58 µm, approximate ratio L/
oral diameter 4.5-7.5.

Measurements of Specimens from Kuwait Waters: Length 214-227 µm; oral diameter 31-40
µm; aboral diameter 16-23 µm; approximate ratio L/oral diameter 3.6-5.0.

Distribution: Eutintinnus macilentus is common at all studied areas of the Arabian Gulf.

a b

Fig. 68 a-c. Variations of lorica shape of Eutintinnus macilentus. Scale – 50 µm.

11 0 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Eutintinnus tenue Kofoid and Campbell, 1929
(Fig. 69 a-c)

Description: Lorica almost cylindrical; with tapering shaft of 2-6 degrees; oral end abruptly
flaring 20-35 degrees into a horizontal brim; aboral end without flare.

Reported Measurements: Length 290 µm (179-238 µm); oral diameter 38-54 µm, approximate
ratio L/oral diameter 4.3-6.1.

Measurements of Specimens from Kuwait Waters: Length 255-265 µm; oral diameter 50-55
µm; aboral diameter 25-26 µm; approximate ratio L/oral diameter 5.0.

Distribution: Eutintinnus tenue is common at all studied areas of the Arabian Gulf.

a b

Fig. 69 a-c. Variations of lorica shape of Eutintinnus tenue. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 11 1


Eutintinnus turgescens (Kofoid and Campbell, 1929)
(Fig. 70 a-d)

Description: Lorica almost cylindrical, 3.2-4.4 oral diameters in length; with subconical shaft of
5-8 degrees, with submedian expansion and strait aboral end from 1/2-2/3 diameter of oral. Oral
end gradually flaring 20-35 degrees to a slightly emergent brim. This species is similar to E. lusus-
undae apart of the median expansion.

Reported Measurements: Length 155-193 µm; oral diameter 39-48 µm, approximate ratio L/
oral diameter 3.2-4.4.

Measurements of Specimens from Kuwait Waters: Length 244 µm; oral diameter 42 µm; aboral
diameter 26 µm; approximate ratio L/oral diameter 5.8.

Distribution: Eutintinnus turgescens is common at all studied areas of the Arabian Gulf.

a
b

c d

Fig. 70 a-d. Variations of lorica shape of Eutintinnus turgescens. Scale – 50 µm.

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Eutintinnus sp.
(Fig. 71 a-d)

Description: Lorica very small, almost cylindrical. Ciliate is bigger in comparison with lorica. Its
body takes all the width of lorica and occupies about 80% of its total length. The wall of lorica
is very thin, transparent and flimsy. In general, most of these tintinnids in the collected samples
have more or less misshapen loricas (Figs. 56 b, d).

Measurements of Specimens from Kuwait Waters: Total length (L) 77-87 µm; oral diameter 15-
20 µm; aboral diameter 12-15 µm; approximate ratio L/oral diameter 3.8-5.3.

Distribution: Eutintinnus sp. was registered near the north coast of Bubiyan Island and in Kuwait
Bay.

a b

c d

Fig. 71 a-d. Variations of lorica shape of Eutintinnus sp. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 11 3


Genus Salpingella Jorgensen, 1924
Reference: Kofoid and Campbell, 1939

Tintinnidae with lorica usually elongated, generally slender, trumpet-shaped, with a short collar
provided with a circular or polygonal oral rim; bowl forming a tubular shaft in the anterior main
part which becomes gradually tapering posteriorly, having a number of raised spiral fins in the
lower region; aboral end typically opening; wall hyaline, composed of a single layer.
Differs from Eutintinnus in having the aboral end contracted, usually to a small terminal cylinder
and in having aboral striae; from Salpingacantha in the absence of teeth in the oral margin;
from Daturella in the rigidity of the lorica and greater aboral taper and contraction; and from
Epicranella in the absence of suboral arches.

Salpingella attenuata Jorgensen, 1924


(Fig. 72 a-c)

Description: Lorica very much elongated, collar is a funnel of 55°-60°, 0.6 oral diameter in length;
bowl cylindrical in the anterior 0.8-0.9 total length, posteriorly a cone (6°-10°) with unevenly
contracted sides; no aboral cylinder; fins 5-7, low blades, 0.2-0.4 total length in length, usually
subvertical or dexiotropic (5°), sometimes giving an appearance of aboral expansion.

Comments: This species differs from S.faurei in the lack of expanded bowl, larger size and fewer
fins, from S. gracilis in size and in having fewer fins, and from S. ricta in more slender proportions
and in the lack of the surface rugose.

Reported Measurements: Length 248-433 µm; oral diameter 32-43 µm, transdiameter of the
bowl 16 µm, approximate ratio L/oral diameter 7.4-15.3.

Distribution: Salpingella attenuata was registered in central and eastern regions of the Arabian
Gulf.

11 4 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a

Fig. 72. Salpingella attenuata. a, b. shape of lorica; c. aboral end.


Stained with Rose Bengal. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 11 5


Salpingella rotundata Jorgensen, 1924
(Fig. 73 a-c)

Description: Lorica small, test-tube-like, inflated posteriorly; bowl cylindrical, with 6-7 low, ridge-
like fins on the posterior 1/3 of the total length; aboral cylinder present.

Comments: This species differs from S. curta in proportions and heavier fins.

Reported Measurements: Length 92-160 µm; oral diameter 13-19 µm, approximate ratio L/oral
diameter 7.0-10.0.

Distribution: Salpingella rotundata was registered in low abundance in the central region of the
Arabian Gulf.

a b

Fig. 73. Salpingella rotundata. a, b. shape of lorica;


c. aboral end. Stained with Rose Bengal. Scale – 50 µm.
c

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Genus Salpingacantha Kofoid and Campbell, 1929
Reference: Kofoid and Campbell, 1929

Tintinnidae with lorica usually elongated, attenuated posteriorly, generally a nail-shaped, posteriorly
truncated, tapering tube; oral margin with blunt or sharp, triangular, flaring, erect or incurved teeth; collar
usually funnel-like or formed only by the swollen base of the teeth; bowl elongated cylindrical or slightly
tapering, usually uniform, sometimes swollen in the middle, posteriorly a narrow, truncated cone; with 5-8
decurrent, low, blade-like fins; antapex with or without a distinct terminal cylinder.
Differs from all other genera of Tintinnidae in the presence of teeth in the oral margin.

Salpingacantha unguiculata Brandt, 1906


(Fig. 74 a-c)

Description: Slender tapering tube, oral region bowl-shaped, but little expanded. Oral margin with 5-8 strong
incurved teeth, margin between flattened and rolled inwards. Shaft sometimes slightly dilated in middle,
contracting to narrow aboral end, with or without aboral cylinder. 7-8 low fins posteriorly.

Reported Measurements: Length 130-290 µm; oral diameter 11-20 µm, approximate ratio L/oral diameter
10.4-14.5.

Distribution: Salpingacantha unguiculata was registered in central and eastern regions of the Arabian Gulf.

a b

Fig. 74 a-c. Variations of lorica shape of Salpingacantha


c unguiculata. Stained with Rose Bengal. Scale – 50 µm.

K u wait I n s titute f or Scienti f ic R e s earc h 11 7


The Geographic Distribution of Tintinnid Species (refer to map on page 105)

Zone I Zone II Zone III Zone IV

Northwestern Central and


Species Waters around
Kuwait Bay region of the Eastern Arabian
Bubiyan Island
(2007-2009) Arabian Gulf Gulf
(2004-2005)
(2003-2009) (Winter 2006)
Leprotintinnus bubiyanicus + + +

Leprotintinnus elongatus + + +

Leprotintinnus nordqvisti + + + +

Tintinnopsis acuminata +

Tintinnopsis ampla + +

Tintinnopsis angusta +

Tintinnopsis baltica + +

Tintinnopsis beroidea + + +

Tintinnopsis compressa +

Tintinnopsis dadayi + +

Tintinnopsis directa + + + +

Tintinnopsis failakkaensis +

Tintinnopsis gracilis + + + +

Tintinnopsis karajacensis + + +

Tintinnopsis lobiancoi + +

Tintinnopsis lohmannii + + +
Tintinnopsis mortensenii + + +
Tintinnopsis nana + + + +

Tintinnopsis orientalis + +

Tintinnopsis parva + + +
Tintinnopsis parvula + + +
Tintinnopsis radix + + + +

Tintinnopsis rotundata +

Tintinnopsis sacculus +

Tintinnopsis schotti + + +

Tintinnopsis tocantinensis + +

Tintinnopsis turbo + +

Tintinnopsis undella +

Tintinnopsis urnula + + +

Tintinnopsis spp. + +

Stenosemella sp. + +

Codonellopsis lusitanica + + +
Codonellopsis morchella + + +

Codonellopsis ostenfeldi + + + +
Dyctiocysta duplex +

Luminella kuwaitensis + + + +
Coxliella annulata + + +

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Zone I Zone II Zone III Zone IV

Northwestern Central and


Species Waters around
Kuwait Bay region of the Eastern Arabian
Bubiyan Island
(2007-2009) Arabian Gulf Gulf
(2004-2005)
(2003-2009) (Winter 2006)
Helicostomella longa +

Metacylis jorgenseni + + +

Metacylis lucasensis +
Metacylis pithos + + + +
Metacylis tropica + + +

Metacylis sp. + +

Ascampbelliella obscura + +

Petalotricha ampulla +

Protorhabdonella curta +

Protorhabdonella simplex +

Rhabdonella conica + +

Rhabdonella striata +

Rhabdonella sp. +

Parundella lohmanni +

Xystonella treforti +

Xystonellopsis gaussi +

Favella adriatica +
Favella campanula + + +

Favella ehrenbergi +

Favella panamensis + + + +
Undella dilatata + +

Amphorellopsis acuta + + +

Amphorides amphora + + +

Amphorides quadrilineata + +

Dadayiella cuspis + +

Dadayiella ganymedes + +

Eutintinnus apertus +

Eutintinnus conicus +

Eutintinnus contractus +

Eutintinnus fraknoi +

Eutintinnus lusus-undae + + +

Eutintinnus macilentus + + +

Eutintinnus tenue + + +

Eutintinnus turgescens + +

Eutintinnus sp. + + +
Salpingella attenuata +

Salpingella rotundata +

Salpingacantha unguiculata +

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1.2. Zoothamnium

Phylum Ciliophora Doflein, 1901


Subphylum Intramacronucleata Lynn, 1996
Class Oligohymenophorea de Puytorac et al., 1974
Subclass Peritrichia Stein, 1859
Order Sessilida Kahl, 1933
Family Zoothamniidae Sommer, 1951
Genus Zoothamnium Bory de St. et al., Vincent, 1826

Reference: Lee et.al., 2000.


(Fig. 75 a-b)

Ciliates, forming branching colonies. Inverted bell-shaped zooids borne upon a contractile
branched stalk which bends in a zigzag fashion (not spirally) by means of a continuous myoneme.
It lies more or less centrally within the stalk and is not sinusoidal when relaxed. Contractile
elements may also extend into the base of the body. Peristomial ciliature extends about one full
turn at unattached end and leads into buccal cavity. Contractile vacuoles empty near base of
buccal cavity. Stressed cells produce basal wreath of cilia and can detach from the stalk and swim
away. Division leads to the production of a basal telotroch, which separates from the parental cell
and swims away. Colonies may be quite large (Fig. 75 a-b).

Fig. 75 a-b. Colony of Zoothamnium sp.

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Flamingos and herons in Shuwaikh Intertidal Zone - Photo by Dr. V. Skryabin - Kuwait Institute for Scientific Research

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1.3. Ellobiopsis

Domain: Eukaryota Whittaker and Margulis, 1978


Kingdom Chromalveolata Adl et al., 2005
Superphylum Alveolata Cavalier-Smith, 1991
Phylum Dinozoa T. Cavalier-Smith, 2004
Infraphylum Protalveolata Cavalier-Smith,1991
Class Ellobiopsea Loeblich, 1970
Order Ellobiopsida Coutiere, 1911
Family Ellobiopsidae Coutiere, 1911
Genus Ellobiopsis Caullery,1910
Ellobiopsis chattoni Caullery, 1910
Reference: Boschma, 1956.

Protista of uncertain position, showing a superficial likeness to peridinians, while in some respects
presenting characters pointing to an affinity with flagellates; on the other hand, a possible affinity
with Fungi (e.g., Saprolegniaceae) is not out of the question. The family consists of heterogeneous
elements: the species of the genus Ellobiocystis are epibiotic, those of the other genera are
parasitic; the species of some genera (Ellobiocystis, Pardobwpsis) do not have an internal stage,
those of other genera (Ellobiopsis, Amallocystis) in all probability are internal parasites before
reaching their full development. Ellobiopsidae are known as epibionts or parasites of pelagic
Crustacea of various groups, and as parasites of non-pelagic Polychaeta (genus Rhizellobiopsis).

Genus Ellobiopsis Caullery, 1910


Body pear-shaped to cylindrical, consisting of a stalk penetrating into the body of the host, a
trophomere, and one or two gonomerea. Parasitic on Copepoda.

Ellobiopsis chattoni Caullery, 1910


Reference: Al-Yamani and Fahmi, 1995
(Fig. 76 a-c)

Body pear-shaped, greatest length about 0.7 mm., greatest thickness about 0.35 mm., the single
gonomere larger than the trophomere. Parasitic on calanoid copepods (Fig. 76).
In Kuwait’s water E. chattoni infected some of the common calanoid copepods such as Acrocalanus
gibber, Paracalanus aculeatus, P. crassirostris and P. parvus as well as their copepodite stages.
Highest incidence of parasitism by E. chattoni was evident during October (Al-Yamani and Fahmi,
1995).

Distribution: Atlantic, Indian and Pacific oceans, Arabian Gulf.

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a

Fig. 76 a-c. Ellobiopsis chattoni on calanoid copepods.

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1.4. Noctiluca

Domain: Eukaryota Whittaker and Margulis, 1978


Kingdom: Chromalveolata Adl et al., 2005
Phylum: Dinoflagellata Bütschli 1885
Superphylum Alveolata Cavalier-Smith, 1991
Class: Noctiluciphyceae Fensome et al., 1993
Order: Noctilucales Haeckel, 1894
Family: Noctilucaceae Saville-Kent, 1881
Genus: Noctiluca Suriray and Lam., 1816

Noctiluca scintillans (Macartney, 1810) Kofoid and Swezy, 1921 = Noctiluca miliaris Suriray, 1836.
Reference: Smithsonian National Museum of Natural History, Internet Site of the Department of Botany, 2009.
(Fig. 77 a-g)

Noctiluca scintillans is a distinctively shaped athecate species in which the cell is not divided into epitheca
and hypotheca. Cells are very large, inflated (balloon-like) and subspherical. The ventral groove is deep and
wide, and houses a flagellum, a tooth and a tentacle. Only one flagellum is present in this species and is
equivalent to the transverse flagellum in other dinoflagellates. The tooth is a specialized extension of the
cell wall. The prominent tentacle is striated and extends posteriorly. Cells have a wide range in size from
200 to 2000 µm in diameter (Fig. 77 a).

N. scintillans is a nonphotosynthetic heterotrophic and phagotrophic dinoflagellate species; chloroplasts


are absent and the cytoplasm is mostly colorless. The presence of photosynthetic symbionts can cause the
cytoplasm to appear pink or green in color. A number of food vacuoles are present within the cytoplasm. A
large eukaryotic nucleus is located near the ventral groove with cytoplasmic strands extending from it to
the edge of the cell. This species is bioluminescent in some parts of the world.
N. scintillans is a cosmopolitan planktonic species common in neritic and coastal regions of temperate,
subtropical and tropical waters. This large phagotrophic species feeding on phytoplankton (mainly diatoms
and other dinoflagellates), protozoans (Fig. 77 b), detritus, copepod’s eggs and nauplii and fish eggs. Big
organisms, as copepodit stages of Copepoda and cyphonautes larvae of Bryozoa are also registered in the
food vacuoles of N. scintillans (Fig. 77 c-d).
Toxic blooms of N. scintillans have been linked to massive fish and marine invertebrate kills. Although this
species does not produce a toxin, it has been found to accumulate toxic levels of ammonia which is then
excreted into the surrounding waters possibly acting as the killing agent in blooms. N. scintillans red tides
frequently form in warm season in many parts of the world, often resulting in a strong pinkish red or orange
discoloration of the water (tomato soup) (Fig. 77 e-g). Green and red blooms often occurm’ Oman sea off
the coast of Oman.

Distribution: Atlantic, Indian and Pacific


oceans, Arabian Gulf.

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b c

d e

Fig. 77. Noctiluca scintillans.


a. general view; b-d. tintinnid Leprotintinnus nordqvisti, Calanoida gen.sp. and cyphonautes larva in food vacuoles
of N. scintillans; e. bloom of N. scintillans in Kuwait water (August 2004); f., g. non-filtered sample of water from the
bloom area.

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2. METAZOA

2.1. Cnidaria

Phylum Cnidaria Verrill, 1865


Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Hydroidomedusae Bouillon et al., 1992
Subclass Anthomedusae Haeckel, 1879
Order Filifera Kuhn, 1913
Suborder Tiarida Haeckel, 1879
Family Pandeidae Haeckel, 1879
Genus Amphinema Haeckel, 1879

Amphinema rugosum (Mayer, 1910)


Reference: Russel, 1955.
(Fig. 78 a-b)

Description: Anthomedusae with considerable apical projection (A in Fig. 78 a, b); two perradial
marginal tentacles (B in Fig. 78 a, b) with basal swellings (c in Fig. 77 a, b). Folded adradial gonads
(D in Fig. 78 a, b) with 3 or 4 folds sloping obliquely downwards toward interradii. 16-24 small
marginal tentaculae. Height up to 6 mm.

Distribution: Indo-Pacifc Region, Atlantic Ocean, Arabian Gulf.

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a

Fig. 78 a-b. Anthomedusae Amphinema rugosum, general view (umbrella height 1.1 mm). Scale - 0.1 mm.
A. apical projection; B. tentacles; C. basal swellings; D. gonad folds.

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Phylum Cnidaria Verrill, 1865
Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Hydroidomedusae Bouillon et al., 1992
Subclass Leptomedusae Haeckel, 1886
Family Aequoreidae Eschscholtz, 1829
Genus Aequorea Peron and Lesueur, 1810

Aequorea pensilis (Eschscholtz, 1829)


References: Bouillon, 1999; Russel, 1970.
(Fig. 79 a-c)

Description: Leptomedusae with lens-shaped central disc (Fig. 79 a). Tentacle bulbs with long
lateral extensions, without excretory papillae (Fig. 79 b, c). Umbrella up to 100 mm wide, more
or less biconvex, with thin margin; manubrium 1/2-2/3 as wide as umbrella; gonads extending
along almost entire length of manubrium; 150-250 radial canals; 10-16 tentacles and as many
small rudimentary bulbs, no excretory papillae but excretory pores present as slits; statocysts
very numerous.

Distribution: Atlantic, Indian Oceans, Arabian Gulf.

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a

b c

Fig. 79. Leptomedusae Aeqourea pensilis (bell diameter 18 mm). a. general view (scale - 1 mm); b., c. tentacle bulb.

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Phylum Cnidaria Verrill, 1865
Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Hydroidomedusae Bouillon et al., 1992
Subclass Leptomedusae Haeckel, 1886
Family Eirenidae Haeckel, 1879
Genus Eirene Eschscholtz, 1829

Eirene viridula (Peron and Lesueur, 1810)


Reference: Bouillon, 1999.
(Fig. 80 a-b)

Description: Umbrella 20-30 mm wide; hemispherical, middle region fairly thick; with slender
gastric peduncle with pyramidal base; manubrium rather small but with 4 long pointed lips
(Fig. 80 a, b), with crenulated margins; with four radial canals; gonads linear, sometimes slightly
sinuous, extending from somewhat beyond base of peduncle to almost bell margin (C in Fig. 80
a, b,); about 70 tentacles of different sizes, large and small frequently alternating (A in Fig. 80 a, b);
bulbs conical with distinct adaxial excretory papillae; 50 or more statocysts.

Distribution: Atlantic, Indian Oceans, Arabian Gulf.

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a b

Fig. 80 a-b. Leptomedusae Eirene viridula (umbrella diameter 10 mm). Scale - 1mm. A. manubrium; B. tentacles; C.
gonad.

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Phylum Cnidaria Verrill, 1865
Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Hydroidomedusae Bouillon et al., 1992
Subclass Leptomedusae Haeckel, 1886
Family Eirenidae Haeckel,1879
Genus Eutima McRady, 1859

Eutima gegenbauri Haeckel


Reference: Russel, 1963.
(Fig. 81 a-c)

Description: Very long prismatic peduncle with broad base (C, D in Fig. 81 a-c); linear gonads
both on peduncle and subumbrellar portions of radial canals (E in Fig. 81 a, b); 8 to 16 or more
marginal tentacles (B in Fig. 81 a, b); 60 to 80 marginal warts; one or two lateral cirri on each side
of tentacles and warts; 8 marginal vesicles each with 6 to 12 or more concretions; up to 20 mm
or more in diameter.

Distribution: Atlantic and Indian Oceans, Arabian Gulf.

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a

Fig. 81. Leptomedusae Eutima gegenbaueri (umbrella diameter 11 mm). a. lateral view (scale - 1 mm); b. apical view;
c. manubrium.
A. manubrium; B. tentacles; C. peduncle; D. base of peduncle; E. gonads.

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Phylum Cnidaria Verrill, 1865
Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Hydroidomedusae Bouillon et al., 1992
Subclass Leptomedusae Haeckel, 1886
Family Phialuciidae Russel, 1953
Genus Octophialucium Kramp, 1925

Octophialucium funerarium (Quoy and Gaimard, 1833)


Reference: Russel, 1963.
(Fig. 82 a-b)

Description: Small stomach without peduncle; normally 8 radial canals (Fig. 82 a, b); no ocelli.
Excretory pores on small adaxial papillae at base of each marginal tentacle; gonads short on
distal ends of radial canals (Fig. 82 a, b); 64 to 128 marginal tentacles; one to three marginal
vesicles between adjacent tentacles, each with one to three concretions; up to 50 mm in
diameter.

Distribution: Indo-Pacific Region, Atlantic Ocean, Arabian Gulf.

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a

Fig. 82 a-b. Leptomedusae Octophialucium funerarium (umbrella diameter 24 mm),


general view. Scale - 1 mm.

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Phylum Cnidaria Verrill, 1865
Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Hydroidomedusae Bouillon et al., 1992
Subclass Leptomedusae Haeckel, 1866
Order Proboscoida Broch, 1910
Family Campanulariidae Jonhston, 1836
Genus Obelia Peron and Lesueur, 1810 (Fig.83)

Obelia spp.
Reference: Bouillon, 1999.
(Fig. 83 a-d)

Description: Campanulariidae with short quadrangular manubrium (A in Fig. 83 a); without


velum; with numerous solid, stiff, not extensile tentacles (B in Fig. 82 a); tentacles with short
endodermal roots extending into bell mesoglea; 8 statocysts located on underside of basal bulbs
of marginal tentacles (C in Fig. 83 a). Umbrella 2.5-6 mm wide, flat, mesoglea very thin; without
gastric peduncle; mouth with 4 simple lips; 4 radial canals (d in Fig. 83 a); gonads spherical to
ovoid, sac-like, hanging from middle to end of radial canals (E in Fig. 83 a). Polyp stage of Obelia
is a component of fowling communities (Fig. 83 c, d).

Distribution: Indo-Pacific Region; Atlantic Ocean; Arabian Gulf.

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a b

c d

Fig. 83. Leptomedusae Obelia spp. (umbrella diameter 1.5 mm). a-b. general view of medusa. (Scale - 0.1 mm);
c. benthic colony of Obelia dichotoma; d. separate zooid of Obelia dichotoma.
A. manubrium; B. tentacles; C. basal bulbs; D. radial canals; E. gonads.

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Phylum Cnidaria Verrill, 1865
Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Hydroidomedusae Bouillon et al., 1992
Subclass Leptomedusae Haeckel, 1866
Family Hydractiniidae Aggassiz, 1862
Genus Podocoryne Lutken, 1850 (Fig.84)

Podocoryne sp.
References: Russell, 1977; Bouillon, 1999.
(Fig. 84 a-d)

Description: Umbrella bell-shaped with rounded apex. Stomach not extending beyond umbrella
margin (Fig. 84 a). Gonads (E in Fig. 84 c) on stomach wall or extending slightly onto radial canals.
Four radial canals (A in Fig. 84 b). Mouth with 4 simple or branching lips with circular nematocyst
clusters (B in Fig. 84 b). 28 marginal tentacles (C in Fig. 84 d) with basal bulbs (D in Fig. 84 d).

Distribution: Indo-Pacific Region; Atlantic Ocean; Arabian Gulf.

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a b

1 mm

c d

Fig. 84. Leptomedusae Podocoryne sp. a - lateral view (umbrella diameter 1.6 mm); b. apical view; c. oral area;
d. margin of umbrella. Scale - 1 mm.
A. radial canals; B. mouth; C. marginal tentacles; D. basal bulbs; E. gonads; F. gastric peduncle.

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Phylum Cnidaria Verrill, 1865
Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Hydroidomedusae Bouillon et al., 1992
Subclass Leptomedusae Haeckel, 1866
Order Proboscoida, Broch, 1910
Family Campanulariidae Jonhston, 1836
Genus Clytia Lamoroux, 1812

Clytia discoida (Mayer, 1900)


Reference: Bouillon, 1999.
(Fig. 85)

Distribution: Leptomedusae with short manubrium; without gastric peduncle; typically with
4 radial canals. Gonads thick, cylindrical, along almost entire length of radial canals (G in Fig.
85). 16 tentacles (T in Fig. 85) . Umbrella 5 mm wide, almost hemispherical; manubrium small,
urn-shaped, with bulging sides (M in Fig. 85); mouth with 4 recurved slightly folded lips; female
gonads with very few large eggs (E in Fig. 85); usually 3 statocysts between adjacent tentacles
(S in Fig. 85).

Distribution: Indo-Pacific Region; Atlantic Ocean; Arabian Gulf.

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Fig. 85. Leptomedusae Clytia discoida (umbrella diameter 5 mm). Scale - 1 mm.
M. manubrium; T. marginal tentacles; G. gonads; E. eggs; S. statocyst.

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Phylum Cnidaria Verrill, 1865
Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Hydroidomedusae Bouillon et al., 1992
Subclass Narcomedusae Haeckel, 1879
Family Aeginidae Gegenbauer, 1857, Maas, 1904
Genus Solmundella Haeckel, 1879

Solmundella bitentaculata Qouy and Gaimard, 1833


Reference: Bouillon, 1999.
(Fig. 86 a-c)

Description: Aeginidae with 8 manubrial pouches (B in Fig. 86 b, c); without peripheral canal
system; 4 peronia (C in Fig. 86 b, c) but only 2 long tentacles (A in Fig. 86 a, b); without secondary
tentacles; without otoporpae. Umbrella up to 12 mm; apical mesoglea very thick; tentacles
issuing from umbrella near apex, 8-32 statocysts.

Distribution: Indo-Pacific Region; Atlantic Ocean; Arabian Gulf.

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a

Fig. 86 a-c. Narcomedusae Solmundella bitentaculata (umbrella diameter 6.1 mm). Scale - 1 mm.
A. tentacles; B. manubrial pouche (lappet); C. peronia.

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Phylum Cnidaria Verrill, 1865
Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Hydroidomedusae Bouillon et al., 1992
Subclass Narcomedusae Haeckel, 1879
Family Cuninidae Bigelow, 1913
Genus Cunina Eschscholtz, 1829

Cunina octonaria McCrady, 1859


Reference: Bouillon, 1999
(Fig. 87 a-b)

Description: Usually 8 (7-9) square manubrial pouches (M in Fig. 87 a), very close together.
Umbrella 5-7 mm wide, flatter than hemispherical; no peripheral canals; thick and broad
ectodermal pad below base of each tentacle; marginal lappets with 2-5 generally 3 statocysts;
otoporae small.

Distribution: Indo-Pacific; Atlantic Ocean; Arabian Gulf.

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a

Fig. 87 a-b. Narcomedusae Cunina octonaria (umbrella diameter 3.5 mm). Scale - 1 mm. General view.
M. manubrial pouches.

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Phylum Cnidaria Verrill, 1865
Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Hydroidomedusae Bouillon et al., 1992
Subclass Trachymedusae Haeckel, 1866
Family Rhopalonematidae Russel, 1953
Genus Aglaura Peron and Lesueur, 1810

Aglaura hemistoma Peron and Lesueur, 1810


Reference: Bouillon, 1999.
(Fig. 88 a-b)

Description: Trachymedusae with slender gastric peduncle (A in Fig. 88 a, b); with 8 sausage-
shaped gonads on peduncle (B in Fig. 88 a, b), not on umbrella; 48-85 tentacles, all alike; umbrella
4-6 mm high, 3-4 mm wide; apex flat; 8 free statocysts.

Distribution: Indo-Pacific Region; Atlantic Ocean; Arabian Gulf.

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a

Fig. 88 a-b. Trachymedusae Aglaura hemistoma (umbrella 1.5 mm height). Scale - 0.1 mm.
A. gastric peduncle; B. gonads.

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Phylum Cnidaria Verrill, 1865
Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Hydroidomedusae Bouillon et al., 1992
Subclass Trachymedusae Haeckel, 1866
Family Geryoniidae Eschscholtz, 1829
Genus Liriope Lesson, 1843

Liriope tetraphyla (Chamisso and Eysenhardt, 1821)


Reference: Bouillon, 1999
(Fig. 89 a-b)

Description: Umbrella 10-30 mm wide, hemispherical; manubrium (A in Fig. 89 a, b) on gastric


peduncle (B in Fig. 89 a, b) of varying size; mouth with 4 lips (C in Fig. 89 b); 1-3 (or more)
centripetal canals in each quadrant; 4 long hollow perradial tentacles (D in Fig. 89, a, b) with
cnidocyst rings and 4 small solid interradial tentacles with adaxial cnidocyst clusters; gonads (E
in Fig. 89 b) variable in shape and size; 8 statocysts.

Distribution: Indo-Pacific Region; Atlantic Ocean; Arabian Gulf.

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a

Fig. 89. Trachymedusae Liriope tetraphyla (umbrella diameter 14 mm. Scale - 1 mm). a. lateral view;
b. apical view.
A. manubrium; B. gastric peduncle; C. lips of mouth; D. perradial tentacles; E. gonads.

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Phylum Cnidaria Verrill, 1865
Class Scyphozoa Goette, 1887
Subclass Scyphomedusae Lankester, 1877
Order Semaeostomeae L. Agassiz, 1862
Family Pelagiidae Gegenbaur, 1856
Genus Sanderia Goette,1886

Sanderia malayensis Goette,1886


References: Uchida and Sugiura,1978; Gershwin and Collins, 2002; Mianzan and Cornelius, 1999.
(Fig. 90 a-d)

Description: Stomach giving rise to 16 radiating pouches which are separate and unbranched (G
in Fig. 90 c); lacking a ring canal; tentacles (B in Fig. 90 b, c) arising from umbrella margin in clefts
between lappets (C in Fig. 90 c), tentacle number - 16; lappets (same sizes) number - 32; mouth
arms long (E in Fig. 90 a, b), pointed, much folded, arms number - 4; rhopalia number – 16 (F in
Fig. 90 c). Gonads not contained in pouches (D in Fig. 90 d).
The sting of most species is painful, and medical attention may be needed.

Distribution: Indo-Pacific Region, Arabian Gulf.

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a b

c d

Fig. 90. Scyphomedusae Sanderia malayensis (bell diameter 53 mm). Scale - 1 cm. a. lateral view; b. aboral view;
c. margin of umbrella; d. oral area.
A. stomach; B. tentacles; C. lappets; D. gonads; E. mouth arms; F. rhopalium; G. stomach pouches; H. gastric cirri.

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Phylum Cnidaria Verrill, 1865
Class Hydrozoa Owen, 1843
Subclass Siphonophorae Eschscholtz, 1829
Order Calycophorae Leuckart, 1854
Family Diphyidae Quoy and Gaimard, 1827
Subfamily Diphyinae Moser, 1825
Genus Diphyes Cuvier, 1817

Diphyes chamissonis (Huxley, 1859)


Reference: Totton, 1954; 1965.
(Fig. 91 a-c)

Description: Anterior nectophore is blunt apically with 5 complete longitudinal ridges (A in


Fig. 91 a). There are 3 prominent ostial teeth (B in Fig. 91 c). Ostial teeth of equal size. Deep
hydroecium (C in Fig. 91 a), only a small basal portion of which may open ventrally. Hydroecium
extends to mid-height of nectophore; with relatively short spindle- shaped somatocyst (D in Fig.
91 a) extending above it. Nectosac with rounded apex lying a short distance from that of the
nectophore. Mouth plate not divided (E in Fig. 91 a, c). Posterior nectophore not developed.

Distribution: Indo-Pacific Region, Arabian Gulf.

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a b

Fig. 91. Siphonophora Diphyes chamissonis (anterior nectophore). Scale - 1 mm.


a-b. general view; c. ostial part.
A. longitudinal ridges; B. ostial teeth; C. hydroecium; D. somatocyst; E. mouth plate.

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2.2. Ctenophora

Phylum Ctenophora Eschscholtz, 1829


Class Tentaculata sensu Mills, 1998
Order Cydippida Gegenbaur, 1856
Family Pleurobranchiidae Chun, 1880
Genus Pleurobrachia Flemming, 1882

Pleurobrachia pileus (O.F.Muller, 1776)


Reference: Mianzan, 1999.
(Fig. 92 a-d)

Description: Egg- to spherical-shaped body; tentacular diameter slightly wider than sagittal;
rows of ciliary combs equal in length (A in Fig. 92 a-c), starting near aboral pole (B in Fig. 92 a-c),
extending more than three quarters of distance toward the mouth (C in Fig. 92 a-c). Tentacle
sheats (D in Fig. 92 a-c) widely separated from stomodeum (E in Fig. 92 a-d). Tentacles with tentilla
simple. Color: comb rows milky opaque, ectomesoderm glassy transparent, tentacle, sheaths and
stomodeum milky or in some specimens dull orange. Length 10-25 mm.

Distribution: Atlantic Ocean, Indo-Pacific Region, Arabian Gulf.

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a b

c d

Fig. 92. Ctenophora Pleurobrachia pileus. a. juvenile (4 mm diameter); b. juvenile (6 mm diameter); c. juvenile (9 mm
diameter); d. juvenile (6 mm diameter). Scale - 1 mm.
A. rows of ciliary combs; B. aboral pole; C. mouth; D. tentacular sheats; E. stomodeum.

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Phylum Ctenophora Eschscholtz, 1829
Class Nuda sensu Mills, 1998
Order Beroida Eschscholtz, 1829
Family Beroidae Eschscholtz, 1829
Genus Beroe Browne, 1756

Beroe sp.
Reference: Mianzan, 1999.
(Fig.93 a-f )

Description: Conical ctenophores with very wide mouth and stomodeum; four meridional
gasrovascular canals (B in Fig. 93 f ) usually with numerous diverticulae (C in Fig. 93 e). Without
tentacles. Body mitten-shaped. Lateral compression very marked. Young B. ovata cannot be
distinguished from B. cucumis. Color dull milky, pink or reddish brown. Height up to 115 mm,
usually 60-70 mm.

Distribution: Atlantic Ocean, Indo-Pacific Region, Arabian Gulf.

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a b

c d

e f

Fig.93. Ctenophora Beroe sp. a. juvenile (height 18 mm); b. juvenile (height 21 mm); c. juvenile (height
25 mm); d. juvenile (height 30 mm); e. wall of the body; f. apical part. Scale - 5 mm.
A. mouth; B. gastrovascular canals; C. diverticulae.

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2.3. Platyhelminthes

Phylum: Platyhelminthes Gegenbaur, 1859


Class: Acoelomorpha Ehlers, 1985
References: Smith and Tyler, 1985; Hooge and Tyler, 2003.
(Fig. 94 a-c)

Description: This group of quite small worms is named acoels because of their lack of a gut
cavity. Their digestive tissue is a syncytium that encompasses food items in large vacuoles rather
than within an epithelially lined lumen as do the guts of other animals. Because no epithelium
delimits the gut contents, they appear solid-bodied (acoel). The digestive tissue in some acoels
is cellular, with spaces between these cells serving as a sort of lumen but not an epithelium-
bounded lumen as in turbellarians. Acoelomorphs resemble other flatworms in many respects,
but have a simpler anatomy. Like flatworms, they have no circulatory or respiratory systems, but
they also lack an excretory system. They have no brain or ganglia, simply a network of nerves
beneath the epidermis, although the nerves are slightly more concentrated toward the forward
end of the animal. The sensory organs include a statocyst, and in some cases, very primitive
pigment-spot ocelli capable of detecting light. They are hermaphrodites, but have no gonads,
and no ducts associated with the female reproductive system. Instead, gametes are produced
from the mesenchymal cells that fill the body between the epidermis and the digestive vacuole.
The cilia of acoels, the locomotory organelles by which they swim, have a unique ultrastructure.
Unlike cilia of other metazoans, the epidermal cilia of acoels are interconnected by a complicated
system of rootlets, and they also have a distinctive shape to their tips--a sort of abrupt narrowing
set off from the main part of the cilium by a sharp shelf. Almost all acoels live in marine habitats,
mostly in between the grains of marine sediments, but some swim in the plankton or creep on
algae and other marine substrates.

Distribution: In Kuwait waters, high concentrations of planktonic acoels are often registered in
the zones of algal blooms.

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a b

Fig. 94 a-c. Variations of body shapes of Acoelomorpha gen.spp.

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Phylum: Platyhelminthes Gegenbaur, 1859
Class: Turbellaria Ehrenberg, 1831
Order: Polycladida Lang, 1884

Müller’s Larvae, Götte’s Larvae


Reference: Brusca and Brusca, 1990.
(Fig. 95 a-b)

Description: In the class Tubellaria, most of species in the order Polycladida produce larvae with
indirect development. In these species, a free-swimming Müller’s larva is formed. This larva is
small, helmet-shaped, and has eight ventrally directed ciliated lobes, which provide propulsion.
Müller’s larva has ciliary bands on swimming arms and apical tuft of cilia on the anterior side.
Mouth is ventral in this larva, and there is no anus. These larvae are pelagic for several days
before they settle to the bottom with their oral surface downward and metamorphose into
juvenile flatworm. Some parasitic polyclades of family Stylochidae produce a free-swimming
four-lobed larvae known as a Götte’s larvae. Some free-living polyclades of family Leptoplanidae
have reduced a four-lobed Müller’s larva, convergently similar to Götte’s larva. Four-lobed larvae
of Turbellaria are displayed in Fig. 95 a, b.

a b

Fig. 95 a-b. Variations of shape of four-lobed Turbellaria Larvae.

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Beach off Khor Al-Sabbiya - Photo by Dr. V. Skryabin - Kuwait Institute for Scientific Research

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2.4. Nemertea (Pilidium Larvae)

Superphylum: Lophotrochozoa (Halanych et al., 1995)


Phylum: Nemertea (Rhyncocoela) Schultze, 1851

Pilidium Larvae
(Fig.96 a-d)

Description: Nemertines are worm- like organisms, the adults of which are generally found
on and in the sea bottom, although some species are deep-water pelagic species. They are
commonly called ribbon-worms, as some are very long, or sometimes proboscis worms, because
they can extend a proboscis to capture their food. Nemertines look like flatworms, but tend to be
larger, thicker bodied, and longer. Size range of adult nemertines is from millimeters to meters.

Their free-swimming, ciliated pilidium larvae are common in plankton samples. The pilidium
larva, which develops from an initial simple ciliated larva has a domed body with a tuft of sensory
cilia sprouting from the top of the apical plate (Fig. 96 a). The lower body is formed by four lobes,
single anterior and posterior lobes that can differ in shape and an identical lateral pair (Fig. 96
d). The edges of the lobes are ciliated (Fig. 96 a) and are used in locomotion and to generate
a current of water carrying food particles toward the mouth, which is situated internal to the
lobes. The epidermis may also contain groups of pigmented cells. Ocelli may be present, but only
develop in species that have them as adults. After a short feeding period the pilidium commences
metamorphosis. The young worm develops within the pilidium, which may continue to feed (Fig.
96 b, c). Metamorphosis is completed when the worm breaks out, often feeding on the remnants
of the pilidium. Size of pilidium larvae is about 0.5-0.8 mm from apical tuft to distal edge of lateral
lobe.

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a b

c d

Fig. 96 a-d. Variations of body shape of Pilidium larvae.


A.c. apical cilia; A.p. apical plate; Ci. cilia; Y.n. developing young nemertine; A.l. anterior lobe; P.l. posterior lobe;
L.l. lateral lobes.

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2.5. Rotifera

Phylum Rotifera Cuvier, 1798


Class Monogononta Plate, 1889
Subclass Eurotatoria De Ridder, 1957
Order Ploima Hudson & Gosse, 1886 (Fig. 97 a-d)
Family Synchaetidae Hudson & Gosse, 1886 (Fig. 97 a-c)
Family Dicranophoridae Harring, 1913 (Fig. 97 d)

Description: Rotifers, or wheel-animalcules, are the smallest of multicellular animals (50-500


μm), with a characteristic crown of fine hairs or cilia surrounding the mouth area. These cilia beat
together in waves to draw water currents (and thus food) toward their mouth. Rotifers primarily
feed on phytoplankton, bacteria, and also detritus. Many rotifers are able to reproduce asexually
when environmental conditions are good, and sexually when environmental conditions are
stressful.

In Kuwait waters, rotifers of the genus Synchaeta (family Synchaetidae) are numerous; rotifers of
the family Dicranophoridae are rarely encountered in the study samples.

Class Monogononta. Males present, but always smaller than females, often dwarfy and with
reduced gut. Most species exhibit heterogony during their life cycle. Body mostly unsegmented,
the foot may exhibit 2-4 ring-like segments. Very diverse morphologies of the rotary organ. Ovary
unpairy. Mainly aquatic (freshwater or marine). Free swimming, semi-sessile or sessile.

Order Ploima. Foot, if present, with 2 toes and 2 glue glands with optional 2 para-glands. Never
completely sessile. Some species are parasitic.

Family Synchaetidae. Sac-, bell- or cone-shaped planktonic species. With either lateral ear-like
ciliar structures on the rotary organ or dorsally and ventrally 2 groups of long, leave- or blade-like
flexible appendages. Feeding is by grabbing-sucking. They are found in freshwater and marine
environments (Fig. 97 a-c).

Family Dicranophoridae. Mostly soft-skinned, longish freshwater and seawater species. Part
of the mastax (chewing stomach) is reorganized to a tong-like grabbing tool. They are found in
freshwater or marine (Fig. 97 d) environments.

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a b

c d

Fig. 97 a-d. Rotifers of Kuwait waters. a-c. Synchaeta spp. d. Dicranophoridae gen.sp. Scale - 0.1 mm.
L. leg; O. ovary; S. stomach.

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2.6. Annelida

Phylum Annelida Lamarck, 1809


Class Polychaeta Grube, 1850
Order Phyllodocida Clark, 1969
Family Lopadorhynchidae Claparede, 1870
Genus Lopadorhynchus Grube, 1875

Lopadorhynchus henseni Reibish, 1893


References: Fernandez-Alamo and Thuesen. 1999; Dales, 1957; Muus, 1953.
(Fig. 98 a-d)

Description: Worms with flattened body (Fig. 98 a). Prostomium broad with four antennae, no
palps and poorly developed eyes (Fig. 98 b). Eversible proboscis, simple and unarmed. With three
tentacular cirri on the first one or two segments which may lack setae. Parapodia uniramous with
dorsal and ventral cirri (Fig. 98 d). Setae mainly compound with serrated end-pieces. Pigidium
without cirri (Fig. 98 c). Two to three pairs of tentacular cirri on the first segment. Next two to
three segments with parapodia modified, with only curved, simple acicular setae. First pairs of
parapodia are stout and shorter than subsequent ones, with simple acicular setae (Fig. 98 b). The
ventral cirri are partially fused to the interior margin of the setigerous lobes (Fig. 98 d).

Distribution: Atlantic Ocean, Indo-Pacific Region, Arabian Gulf.

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0.1 mm

0.1 mm

1 mm

Fig. 98. Polychaeta Lopadorhynchus henseni. a. general view (scale - 1 mm); b. oral end (scale - 0.1 mm);
c. pygidium (scale - 0.1 mm); d. parapodia with acicular setae.

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Phylum Annelida Lamarck, 1809
Class Polychaeta Grube, 1850
Order Phyllodocida Clark, 1969
Family Lopadorhynchidae Claparede, 1870
Genus Pelagobia Greef, 1879

Pelagobia longicirrata Greef, 1879


References: Fernandez-Alamo and Thuesen. 1999; Dales, 1957; Muus, 1953.
(Fig. 99 a-c)

Description: Prostomium triangular (A in Fig. 99 b) with four small antennae and a pair of small
eyes (Fig. 99 a, b). Tentacular segment with a few setae and two long tapered cirri (C in Fig. 99 b).
Later parapodia with a conical setigerous lobe and longer dorsal and ventral cirri (D in Fig. 99 c),
essentially similar, more or less cylindrical. Compound setae with basal joints are smooth.

Distribution: Atlantic Ocean, Indo-Pacific Region, Arabian Gulf.

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a

Fig. 99. Polychaeta Pelagobia longicirrata (length 17 mm). a. general view (scale - 1 mm); b. anterior part;
c. posterior part.
A. prostomium, B. antennae; C. tentacular cirri; D. dorsal and ventral cirri.

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2.7. Brachiopoda Larvae

Phylum Brachiopoda Duméril, 1806


Class Lingulata Gorjansky and Popov, 1985 (Fig. 100 a-c)
References: Pennington and Stricker, 2002; Brusca, and Brusca, 1990.

Description: Brachiopods or ‘lamp shells’ are sedentary filter-feeding invertebrates that


superficially resemble the bivalved mollusks. However, basic features of their anatomy (body
axes, gills, foot, gonads) are fundamentally different from those found in bivalves. Brachiopods
have often been considered to be evolutionarily unrelated to the Mollusca. Brachiopods come
in two easily distinguished varieties. Articulate brachiopods have a hinge-like connection or
articulation between the shells; whereas, inarticulate brachiopods are not hinged and are held
together entirely by musculature.

The larvae of inarticulates swim as plankton for months and are like miniature adults, with valves,
mantle lobes, a pedicle that coils in the mantle cavity, and a small lophophore, which is used for
both feeding and swimming (Fig. 100, a, b, c).

The main features of the Lingulata are dorsal and ventral shell valves underlain by mantle tissue,
a complete gut, a developing pedicle, and a lophophore. Each shell valve is divided into a
posterior semicircular embryonic shell or protegulum, which develops before a larva hatches
from its egg membrane, and a larger, circular to ovoid larval shell, which is added as larvae grow
in the plankton (in discinacean, larvae protegulum is absent). The lophophore consists of pairs of
cirri that bud off sequentially on either side of the median tentacle, so that the youngest tentacle
pair is most anterior. The median tentacle is a putative sensory organ characteristic of lingulacean
and discinacean larvae. The anterior lip of the mouth is termed the epistome.

Distribution: Arabian Gulf and other seas.

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a
b

Fig. 100. a-c. Brachiopod’s larvae.


C. cirri; E. epistome; M. mouth; S. stomach; SV. shell valves; LS. larval setae; PLS. principal larval setae.

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2.8. Mollusca

Phylum Mollusca Linnaeus, 1758


Class Gastropoda Cuvier, 1797
Subclass Prosobranchia Thiele, 1921
Order Mesogastropoda Thiele, 1921
Superfamily Heteropoda Lamarck , 1812
Family Atlantidae Rang, 1829
Genus Atlanta Lesueur, 1817

Atlanta sp. (Fig.101 a-b)

Description: Dextrally (right-handed) coiled small transparent shell (Fig. 101 a, b). Shell surface
smooth. The last body whorl is greatly enlarged and laterally flattened, bearing a keel along its
outer edge (B in Fig. 100 b). When the soft parts of mollusk completely retracts into the shell (A
in Fig. 101 a), almost one half of the enlarged, outermost shell whorl remains unoccupied. Larval
shell is present as the inner, helical part of the adult shell. Number of spire whorls is 2 (Fig. 101
a, b).

It is to be noted that four morphological characters (shell, eyes, operculum, and radula) are of
great importance in the identification of heteropods, particularly Atlantidae.

Distribution: Arabian Gulf and other seas.

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a

0.1 mm

Fig. 101 a-b. Heteropoda Atlanta sp. juv.


A. soft parts of body; B. keel. Scale - 0.1 mm.

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Phylum Mollusca Linnaeus, 1758
Class Gastropoda Cuvier, 1797
Subclass Euthyneura (=Opisthobranchia) Spengel, 1881
Order Thecosomata de Blainville, 1824
Suborder Euthecosomata Meisenheimer, 1905
Family Limacinidae Grey, 1847
Genus Limacina, Bosc 1817

Limacina bulimoides (d’Orbigny, 1834)


(Fig. 102)

Description: Spire with 5 whorls. Thicker parts of the shell near aperture brownish. Shell
transparent. Shallow suture light-brown. The outer surface of the shell is smooth. Small rostrum
more pointed in older than in young specimens. Shell height 1.3 mm, maximum diameter 0.6
mm (Fig. 102).

Distribution: Tropical and subtrorieal regious of Atlantic, Indian and Pacific oceans, Arabian Gulf.

Fig. 102. Limacina bulimoides. Scale - 0.1 mm.

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Phylum Mollusca Linnaeus, 1758
Class Gastropoda Cuvier, 1797
Subclass Euthyneura (=Opisthobranchia) Spengel, 1881
Order Thecosomata de Blainville, 1824
Suborder Euthecosomata Meisenheimer, 1905
Family Cavoliniidae Fisher, 1883
Subfamily Clioinae van der Spoel, 1972
Genus Creseis Rang, 1828

Creseis chierchiae (Boas, 1886)


(Fig. 103 a-d)

Description: Shell transparent (Fig. 103 a, b), cross-section circular, surface with conspicuous
transverse rings (A in Fig. 103 a, b), with wing protrusions. Embryonic shell (B in Fig. 103 a, b) with
anteriorly convex, posteriorly concave sides, blunt tip. Clear constriction separates embryonic
from adult shells. Protoconch is conical. Shell length 1.6 mm (exceptionally up to 2.7 mm), width:
0.2 mm, embryonic shell 0.4-0.5 mm long. It is encountered in the Arabian Gulf.

a b

c d

Fig. 103. Pteropod Creseis chierchiae. a. general view (shell length 1.55 mm); b. general view (shell length 1.39 mm);
c. early juvenile stage (stained with Rose Bengal), d. juvenile stage. Scale - 0.1 mm.
A. transverse rings; B. embryonic shell.

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Phylum Mollusca Linnaeus, 1758
Class Gastropoda Cuvier, 1797
Subclass Euthyneura (=Opisthobranchia) Spengel, 1881
Order Gymnostomata de Blainville, 1824
Suborder Gymnostomata de Blainville, 1824
Family Pneumodermatidae Pelseneer, 1887 (Fig. 104 a-b)

Description: Organism with either two lateral sucker arms, or one median and two lateral arms.
Buccal mass with well-developed radula and pair of hook sacs filled with large hooks. Gills usually
at posterior body pole and in middle of right lateral side. Penis, absent during female stage, is
evaginable and in some species provided with an extra sucker, in species without sucker a special
median tubercle appears at base of posterior footlobe. Skin usually with chromatophores; with
acetabuliferous arms. With at least one gill (Fig.104).

Distribution: Arabian Gulf and other seas.

a b

Fig. 104. a-b. Naked pteropoda Pneumodermatidae gen.sp.

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Phylum Mollusca Linnaeus, 1758
Class Cephalopoda Cuvier, 1797
Subclass Coleoidea Bather, 1888
Superorder Decapodiformes Young, et al., 1998
Order Sepiolida Fioroni, 1981
Family Sepiolidae Leach, 1817 (Fig.105)

Reference: Jereb and Roper, 2005.

Bobtail squids (order Sepiolida) are a group of cephalopods closely related to cuttlefish.

Description: Mantle of bobtail squids of family Sepiolidae is broad, oval; posterior mantle
margin rounded. Dorsal mantle fused to head, or free from head. Mantle cavity divided by thin
septum. Fins wide; rounded, semicircular, or kidney-shaped, with pronounced anterior lobes, or
“earlets”; fin attachment short, fin length exceeds attachment length (F in Fig. 105). Large eyes
covered by corneal membranes. Mantle-locking cartilage simple, linear. Arms short; protective
membranes absent in both sexes. Tentacles retractable, each bears a well defined club. Internal
gladius present, rudimentary, chitinous, or absent. Mantle length of adults up to 80 mm, rarely
100 mm. Eggs large, development direct.

Distribution: Arabian Gulf.

Fig. 105. Immature bobtail squid Sepiolidae gen.sp. Scale - 0.1 mm.
F. fin.

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2.9. Lophotrochozoa Larvae

Superphylum Lophotrochozoa (Halanych et al., 1995)


Phylum Phoronida Hatschek, 1888
Class Phorona Hatschek, 1888
Order Phoronida Wright, 1856
Family Phoronidae Wright, 1856

Actinotrocha Larvae
Reference: Emig, 1982.
(Fig. 106 a-d)

Description: The actinotroch larva of Phoronida, bears an anterior preoral lobe on which is
located the nervous ganglion (on the apical area), a tentacular ridge, a pair of protonephridia,
and posteriorly a ciliated ring around the anus (Fig. 106 d). The larvae undergo a planktotrophic
development from 2–3 weeks and settle after about 20 days. Metamorphosis is very fast,
occurring in less than 30 minutes and leading to a slender young phoronid.

Distribution: Arabian Gulf and other seas.

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a b

c d

Fig. 106. a-d. Different age stages of Actinotroch larvae.


P.L. preoral lobe; T. tentacles; Tr. trunk; S. stomach; I. intestine; A. anus.

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Superphylum Lophotrochozoa (Halanych et al., 1995)
Phylum Bryozoa (Ectoprocta) (Ehrenberg, 1831)
Class Gymnolaemata Allman, 1856
Order Cheilostomata Levinsen, 1906

Cyphonautes Larvae
Reference: Todd et al., 1990.
(Fig.107 a-d)

Description: The adults of Bryozoa are colonial, forming gelatinous or firm encrusting mats, or
raised coral - like structures (Fig. 107 c, d). Bryozoan larvae are 0.2-0.7 mm in length, vary in form,
but all have a band of cilia round the body which enables them to swim, a tuft of cilia at the top,
and an adhesive sac that everts and anchors them when they settle on a surface. Some species
of class Gymnolaemata produce cyphonautes larvae which have little yolk but a well-developed
mouth and gut, and live as plankton for a considerable time before settling. These larvae have
triangular shells of chitin, with one corner at the top and the base open, forming a hood around
the downward-facing mouth (Fig. 107 a, b).

Distribution: Arabian Gulf and other seas.

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a b

c d

Fig. 107. a-b. Cyphonautes larvae; c. general view of colony of Bryozoa Bugula neritina; d. polyps of B.neritina.
A. apical sense organ; Ci. band of cilia.

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2.10. Cirripedia

Phylum Arthropoda Latreille, 1829


Subphylum Crustacea Brünnich, 1772
Class Maxillopoda Dahl, 1956
Subclass Thecostraca Gruvel, 1905
Infraclass Cirripedia Burmeister, 1834
Order Sessilia Lamarck, 1818
Suborder Balanomorpha Pilsbry, 1916
Genus Balanus Da Costa, 1778

Balanus amphitrite Darwin, 1854


Reference: Nicholson, 1880.
(Fig. 108 a-d)

Description: Barnacles are encrusters, attaching themselves permanently to a hard substrate.


The most common, “acorn barnacles” (Order Sessilia) are sessile, growing their shells directly onto
the substrate (Fig. 108 d).

The nauplius stage of barnacles has an unsegmented, pyriform body, enclosed in a carapace,
provided with long caudal and dorsal spines (Fig. 108 a). There are three pairs of limbs, of which
the first pair (representing the antennas) is undivided, while the two hinder pairs are bifid, and all
carry natatory bristles. Very large labrum placed in front of the mouth, well-developed alimentary
tube terminates by a distinct anus at the root of the caudal spine. There is at first merely a simple
central eye; but in the late nauplius, two compound lateral eyes are developed in addition.

The nauplius passes into its second condition or cypris-stage. It is enclosed in an oval, bivalved,
mussel-shaped shell, with an opening along the ventral margin (Fig. 108 b,c). The second and third
pairs of the appendages of the nauplius disappear, and the first pair of appendages constitutes
strong four-jointed antennae, the last segment of which is disc-shaped, and is pierced centrally
by a pore, which is the opening of the excretory duct of the cement-glands, these organs being
situated at the bases of the antennae. The thorax has developed upon its sides six pairs of forked
natatory limbs; the abdomen is rudimentary, three-jointed, with terminal forked swimming
appendages. The cypris larva does not feed, but is nourished by means of accumulation of fatty
matter, which had been stored up by the nauplius in the cephalic and dorsal regions of the body.

The cypris larva is the final larval stage of barnacles. Metamorphosis into a cyprid usually follows
5 or 6 planktotrophic nauplius stages and the time that the cypris of Balanus amphitrite spends
in the plankton is a few days. During this part of the life cycle, the cypris searches for a place to
settle. It explores surfaces with modified antennules; once the cypris has found a potentially
suitable area, it attaches using its attachment antenna and a secreted glycoproteinous substance.

Distribution: Widely distributed in tropical and warm temperate seas.

18 2 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

c d

Fig. 108. a. Nauplius of Cirripedia; b-c. cypris larvae of Balanus amphitrite; d. adult stage of B.amphitrite.
A.a. attachment antenna.

K u wait I n s titute f or Scienti f ic R e s earc h 18 3


2.11. Cladocera

Phylum Arthropoda Latreille, 1829


Subphylum Crustacea Brunnich,1772
Class Branchiopoda Latreille,1817
Subclass Phyllopoda Preuss,1951
Order Diplostraca Gerstaecker,1866
Suborder Cladocera Latreille,1829
Infraorder Onychopoda Sars,1865
Family Sididae Baird, 1850
Genus Penilia Dana, 1849

Penilia avirostris Dana, 1852


(Fig. 109 a-c)

Description: Body transparent; rostrum sharply pointed in female (A in Fig. 109 a), rounded in
male. Antennules short with several distal aesthetascs (small sensory setae covered by delicate
cuticle, often projecting from antennule, also known as olfactory hairs) in females, but elongated
and almost reaching posterior end of carapace in males. Antennae consisting of a large basipod
(B in Fig. 109 a), and 2-segmented exo- and endopods (C, D in Fig. 109 a). Exopods bearing 2
swimming setae on first segment and 6 on the second, endopods with 1 (on first segment) and
4 (on the second) swimming setae (setal formula: 2,6/1,4). Carapace with a prominent spine on
posterior end (E in Fig. 109 a), with row of minute spines on both ventral and postero-dorsal
edges, and much finer spinules between these spines. Postabdomen large, ending in a pair of
long terminal claws (F in Fig. 109 a) with 2 spines (distal one much longer than proximal) in
postero-basal portion. Dorso-posterior corner of carapace depressed, bearing a pair of long
caudal setae (G in Fig. 109 a). Six pairs of thoracic limbs (thoracic legs, or thoracopods) (H in
Fig. 109 a), last one short and reduced; all limbs biramous, endopod indistinctly segmented
with many minute plumose setae directing posteriorly, exopod consisting of a flat lamella with
several thick plumose setae. Endopod of male thoracic limb 1 with a sharp recurved claw on
distal end (terminal hook). In males a pair of copulatory appendages (penes) located posterior to
thoracic limb 6. Resting eggs: usually produced by larger females in population; grayish, ovoid
and somewhat flattened.
Total length, females: 0.4-1.2 mm; males: 0.7-0.9 mm.

Distribution: Widely distributed in tropical and warm temperate seas.

18 4 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

Fig. 109. Penilia avirostris. a, b. left lateral view: c. dorsal view. Scale - 0.1 mm.
A. rostrum; B. basiopod of antennae; C. exopod of antennae; D. endopod of antennae; E. carapace spines;
F. postabdomen with terminal claws; G. caudal setae; H. thoracopods.

K u wait I n s titute f or Scienti f ic R e s earc h 18 5


Phylum Arthropoda Latreille, 1829
Subphylum Crustacea Brunnich,1772
Class Branchiopoda Latreille,1817
Subclass Phyllopoda Preuss,1951
Order Diplostraca Gerstaecker,1866
Suborder Cladocera Latreille,1829
Infraorder Onychopoda Sars,1865
Family Podonidae Mordukhai-Boltovskoi, 1968
Genus Pleopis Dana, 1852

Pleopis polyphemoides (Leuckart, 1859) [=Podon polyphemoides (Leuckart,


1859)]
(Fig. 110 a-c)

Description: Body small. Brood chamber varying from hemi-spherical to spherical (A in Fig. 110
a). Caudal furca short, in outline almost an equilateral triangle (B in Fig. 110 a). Antennae (C in
Fig. 110 a): exopod segments 1-4 with 0,1,2,4 swimming setae. Exopods of thoracic limbs 1-4
with 3,3,3,2 setae, respectively. Resting eggs: small, spherical, 0.15-0.21 mm in diameter (mean
0.18 mm); greenish-brown; numerous small pores scattered throughout outer egg membrane;
perimeter obscured with the ornamentation of membrane.

Total length (standard length) 0.3-0.7 mm (0.2-0.6 mm) in female, 0.4-0.7 mm (0.4.-0.7 mm) in
male.

Distribution: Widely distributed in tropical and warm temperate seas.

18 6 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a b

0.1 mm

Fig. 110. a-c. Pleopis polyphemoides, right lateral view. Scale - 0.1 mm.
A. brood chamber; B. caudal furca; C. antennae.

K u wait I n s titute f or Scienti f ic R e s earc h 18 7


Phylum Arthropoda Latreille, 1829
Subphylum Crustacea Brunnich,1772
Class Branchiopoda Latreille,1817
Subclass Phyllopoda Preuss,1951
Order Diplostraca Gerstaecker,1866
Suborder Cladocera Latreille,1829
Infraorder Onychopoda Sars,1865
Family Podonidae Mordukhai-Boltovskoi, 1968
Genus Pseudevadne Claus, 1877

Pseudevadne tergestina (Claus, 1877) (=Evadne tergestina Claus, 1877)


(Fig. 111 a-c)

Description: Body oval. Carapace always with rounded posterior end, lacking spine (Fig. 111);
varying in shape from elongate-oval to hemispherical. Elevator muscles of antenna diverging
dorsally. Exopods of thoracic limbs 1-4 with 2,3,3,1 setae, respectively. Several rows of hypodermal
cells (C in Fig. 111 a) present in brood pouch (often disappearing in preserved specimens).

Total length (standard length) in female 0.3-1.3 mm (0.3-0.9 mm); in male 0.5-0.8 mm (0.3-0.6
mm).

Distribution: Widely distributed in tropical and warm temperate seas.

18 8 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


a

Fig. 111. a-c. Pseudevadne tergestina, left lateral view. Scale - 0.1 mm.
A. antenna; B. posterior end of carapace; C. hypodermal cells.

K u wait I n s titute f or Scienti f ic R e s earc h 18 9


2.12. Ostracoda

Phylum Arthropoda
Subphylum Crustacea Brunnich,1772
Class Ostracoda Latreille,1802
Subclass Myodocopa Sars, 1866
Order Halocyprida Dana,1853
Suborder Halocypridina Dana,1853
Superfamily Halocypridoidea Dana,1853
Family Halocyprididae Dana,1853
Subfamily Euconchoecinae
Genus Euconchoecia Muller, 1890

Euconchoecia aculeata Scott, 1894


(Fig. 112)

Description: The shell is delicate and elongate; the height of the male and female shells is clearly
less than half the length (Fig. 112). There are spines or points at both postero-dorsal corners in
both sexes (F in Fig. 112) the one on the right longer than the one on the left shell. The postero-
ventral and antero-ventral corners are rounded (A and B Fig. in 112). The asymmetrical glands
open on both sides on the posterior margin, just below the postero-dorsal corner (G in Fig. 112).
No other glands have been observed. The rostrum (C in Fig. 112) has small points on both sides,
the one on the left longer than the one on the right. The frontal organs of both sexes are slim
and undifferentiated. The male and female first antennae have over 20 sensory filaments. In
Euconchoecia the shape of the endopod of the second antenna is quite different from Conchoecia
and the processus mammilaris is lacking. The longest seta of the endopod of the male (not
female) second antenna is considerably longer than the shell (D in Fig. 112). The shape of the
penis is distinctive. The furca has seven claws (E in Fig. 112). The female may carry two to eight
large eggs, up to 0.25 mm diameter, between the back of the body and the shell.
Length. Females 1.1-1.5 mm, males 1.15-1.25 mm.

Distribution: Atlantic Ocean, Indo-Pacific Region, Arabian Gulf.

19 0 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Fig. 112. Euconchoecia aculeata (female). Scale - 0.1 mm.
A. antero-ventral corner of carapace; B. postero-ventral corner of carapace; C. rostrum; D. seta of
second antenna endopod; E. claws of furca glands; F. spine; G. glands.

K u wait I n s titute f or Scienti f ic R e s earc h 19 1


Phylum Arthropoda
Subphylum Crustacea Brunnich,1772
Class Ostracoda Latreille,1802
Subclass Myodocopa Sars, 1866
Order Myodocopida Sars, 1866
Suborder Myodocopina Sars, 1866
Family Cypridinidae Baird, 1850
Subfamily Cypridininae Baird, 1850
Genus Cypridina H. Milne Edwards, 1840

Cypridina sp.
(Fig. 113 a-g)

Description: The genus Cypridina has a number of distinguishing characters, e.g. shell-form and
upper lip, well separated from the other genera of Cypridininae. The species are very similar
in structure. The main distinguishing marks are found in the shell (its form and especially its
differing armature of spines and bristles) and in the furca. The shell is rather elongate, the height
being between 50-60% of the length. In the females it is a little higher than in the males (Fig. 113).
The antero-superior corner of rostrum is protruding over the front margin; the “corner” is as a rule
rounded, in a single species pointed (Fig. 113 b). The incisur is rather short and broad (B in Fig.
113 b). The shell margin below the incisur is in few species bare, in the remainder it has a row of
hairs varying between 1-2 and 27; the numbers of these hairs offer good specific distinguishing
marks (D in Fig. 113 b). The posterior shell process is always well-developed, varying somewhat
in size and form (E in Fig. 113 a, c). The ventral shell-list in front of the posterior process either
almost follows the contour of the process or it is straight or even slightly concave (Fig. 113 c); its
edge is in some species bare, in others only the right or both ridges are provided with a number
of spines; a ridge with spines is always more straight and stronger chitinized than one without
spines. In some species spines are found on the medial surfaces of the process. Medially across
the rostrum is a varying, but always small, number of rather weak bristles as a rule are arranged
in an irregular, vertical row behind the anterior margin and along a short, low ridge which turns
backward and upward from the pointed tip of the rostrum (F in Fig. 113 b). At the bottom of the
incisur is the usual pair of spine-like bristles (C in Fig. 113 b). In some species the margin off the
lower rounding of the incisur bears 6-7 backwards pointing spines. The margin of the shell just
below the incisur can be smooth, dentated or serrated. The ventral list is provided with a few
widely spread, weak bristles.

Distribution: Arabian Gulf and othe seas.

19 2 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


b a

g
d e f

Fig. 113. Cypridina sp. (female). a. general view; b. anterior part of carapace (left shell); c. ventral shell-list; d. first
antenna; e. mandible; f. furca with claws; g. 6th limb. Scale - 0.1 mm.
A. rostrum; B. incisur; C. bristles; D. hairs; E. posterior process; F. rostrum bristles.

K u wait I n s titute f or Scienti f ic R e s earc h 19 3


Index of Scientific Names
Acanthephyra sp. Vol.2, 146 Clytemnestra scutellata Vol.2, 125
Acartiidae Vol.2, 80 Clytia discoida Vol.1, 140-141
Acartia Vol.2, 80 Codonellidae Vol.1, 20
Acartia (Acanthacartia) fossae Vol.2, 82-83 Codonellopsidae Vol.1, 56
Acartia (Acartiella) faoensis Vol.2, 88-89 Codonellopsis Vol.1, 56
Acartia (Odontacartia) amboinensis Vol.2, 84-85 Codonellopsis lusitanica Vol.1, 56-57
Acartia (Odontacartia) ohtsukai Vol.2, 86-87 Codonellopsis morchella Vol.1, 58
Actinotrocha Larvae Vol.1, 178-179 Codonellopsis ostenfeldi Vol.1, 59
Aequorea pensilis Vol.1, 128-129 Copepoda Vol.2, 1-7
Aglaura hemistoma Vol.1, 146-147 Copilia Vol.2, 108
Alpheus spp. Vol.2, 149 Copilia mirabilis Vol.2, 108-109
Amphinema rugosum Vol.1, 126-127 Corycaeidae Vol.2, 112
Amphorellopsis Vol.1, 96 Corycaeus Vol.2, 112
Amphorellopsis acuta Vol.1, 96-97 Corycaeus (Dithrichocorycaeus) andrewsi Vol.2, 113
Amphorides Vol.1, 98 Corycaeus (Dithrichocorycaeus) dahli Vol.2, 114-115
Amphorides amphora Vol.1, 98-99 Corycaeus (Dithrichocorycaeus) lubbocki Vol.2, 116
Amphorides quadrilineata Vol.1, 100 Corycaeus (Onychocorycaeus) agilis Vol.2, 117
Appendicularia sicula Vol.2, 174 Corycaeus (Onychocorycaeus) pacificus Vol.2, 118
Ascampbelliedidae Vol.1, 76 Coxliella Vol.1, 64
Ascampbelliella Vol.1, 76 Coxliella annulata Vol.1, 64-65
Ascampbelliella obscura Vol.1, 76-77 Creseis chierchiae Vol.1, 175
Atlanta sp. Vol.1, 172-173 Cunina octonaria Vol.1, 144-145
Cyclopoida Vol.2, 94-95
Balanus amphitrite Vol.1, 182-183 Cyphonautes Larvae Vol.1, 180-181
Beroe sp. Vol.1, 156-157 Cypridina sp. Vol.1, 192-193
Bestiolina Vol.2, 20
Bestiolina arabica Vol.2, 20-21 Dadayiella Vol.1, 101
Brachiopoda Larvae Vol.1, 170-171 Dadayiella cuspis Vol.1, 101
Branchiostoma sp. Vol.2, 180-181 Dadayiella ganymedes Vol.1, 102
Dardanus sp. Vol.2, 161
Calanidae Vol.2, 10 Dicranophoridae Vol.1, 164-165
Calanoida Vol.2, 10 Dictyocystidae Vol.1, 60
Calanopia Vol.2, 58 Dictyocysta Vol.1, 60
Calanopia elliptica Vol.2, 58-59 Dictyocysta duplex Vol.1, 60-61
Calanopia minor Vol.2, 60-61 Diogenes sp. Vol.2, 160
Callianassa sp. Vol.2, 152 Diphyes chamissonis Vol.1, 152-153
Candaciidae Vol.2, 56 Dithrichocorycaeus Vol.2, 113
Candacia Vol.2, 56
Candacia bradyi Vol.2, 57 Ebalia sp. Vol.2, 164
Canthocalanus Vol.2, 12 Ectinosomatidae Vol.2, 120
Canthocalanus pauper Vol.2, 12-13 Eirene viridula Vol.1, 130-131
Catostylus mosaicus Vol.2, 182-183 Ellobiopsis chattoni Vol.1, 122-123
Centropagidae Vol.2, 42 Eucalanidae Vol.2, 29
Centropages Vol.2, 42 Euchaetidae Vol.2, 36
Centropages furcatus Vol.2, 44-45 Euchaeta Vol.2, 36
Centropages orsinii Vol.2, 46-47 Euchaeta concinna Vol.2, 38-39
Centropages tenuiremis Vol.2, 47 Euchaeta rimana Vol.2, 40-41
Cirripedia Vol.1, 182-183 Euterpinidae Vol.2, 124
Clausocalanidae Vol.2, 33 Euterpina Vol.2, 124
Clausocalanus Vol.2, 33 Euterpina acutifrons Vol.2, 124
Clausocalanus minor Vol.2, 34-35 Eutima gegenbauri Vol.1, 132-133
Clytemnestridae Vol.2, 124 Euconchoecia aculeata Vol.1, 190-191
Clytemnestra Vol.2, 125 Eutintinnus Vol.1, 103

19 4 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


Eutintinnus apertus Vol.1, 103 Metacylis jorgenseni Vol.1, 68-69
Eutintinnus conicus Vol.1, 104 Metacylis lucasensis Vol.1, 70-71
Eutintinnus contractus Vol.1, 106-107 Metacylis pithos Vol.1, 72-73
Eutintinnus fraknoi Vol.1, 108 Metacylis sp. Vol.1, 75
Eutintinnus lusus-undae Vol.1, 109 Metacylis tropica Vol.1, 74
Eutintinnus macilentus Vol.1, 110 Metapenaeus ensis Vol.2, 128
Eutintinnus sp. Vol.1, 113 Microsetella Vol.2, 120
Eutintinnus tenue Vol.1, 111 Microsetella sp. Vol.2, 120-121
Eutintinnus turgescens Vol.1, 112 Miraciidae Vol.2, 122

Favella Vol.1, 89 Noctiluca scintillans Vol.1, 124-125


Favella adriatica Vol.1, 89
Favella campanula Vol.1, 90 Obelia spp. Vol.1, 136-137
Favella ehrenbergi Vol.1, 91 Octophialucium funerarium Vol.1, 134-135
Favella panamensis Vol.1, 92-93 Odontacartia Vol.2, 84
Fritillaria pellucida Vol.2, 184-185 Oikopleura dioica Vol.2, 173
Fritillaria sp. Vol.2, 175 Oithonidae Vol.2, 94
Oithona Vol.2, 94-96
Galathea sp. Vol.2, 158 Oithona attenuata Vol.2, 96-97
Oithona brevicornis Vol.2, 98-99
Harpacticoida Vol.2, 120 Oithona nana Vol.2, 98-99
Helicostomella Vol.1, 66 Oithona plumifera Vol.2, 100
Helicostomella longa Vol.1, 66-67 Oithona sp. Vol.2, 102-103
Oncaeidae Vol.2, 104
Ilyoplax frater Vol.2, 168 Oncaea Vol.2, 104
Oncaea clevei Vol.2, 106-107
Labidocera Vol.2, 62 Onychocorycaeus Vol.2, 116
Labidocera acuta Vol.2, 62-65
Labidocera bengalensis Vol.2, 66-67 Pachycheles sp. Vol.2, 159
Labidocera kroyeri Vol.2, 58-69 Paguristes sp. Vol.2, 162
Labidocera minuta Vol.2, 70-71 Pagurus sp. Vol.2, 163
Labidocera sp. Vol.2, 72-73 Palaemon sp. Vol.2, 147
Laomedia sp. Vol.2, 153 Paracalanidae Vol.2, 14-15
Latreutes sp. Vol.2, 150 Paracalanus Vol.2, 22
Leprotintinnus Vol.1, 14 Paracalanus indicus Vol.2, 22-23
Leprotintinnus bubiyanicus Vol.1, 14-15 Paracalanus sp. Vol.2, 24-25
Leprotintinnus elongatus Vol.1, 16-17 Parapenaeopsis stylifera Vol.2, 129
Leprotintinnus nordqvisti Vol.1, 18-19 Parthenope sp. Vol.2, 166
Leucosia sp. Vol.2, 165 Parundella Vol.1, 86
Limacina bulimoides Vol.1, 174 Parundella lohmanni Vol.1, 86
Liriope tetraphyla Vol.1, 148-149 Parvocalanus Vol.2, 26
Lopadorhynchus henseni Vol.1, 166-167 Parvocalanus crassirostris Vol.2, 26-27
Lucifer hanseni Vol.2, 135-137 Parvocalanus elegans Vol.2, 28
Lucifer typus Vol.2, 138 Pelagobia longicirrata Vol.1, 168-169
Luminella Vol.1, 62 Penaeus semisulcatus Vol.2, 130-131
Luminella kuwaitensis Vol.1, 62-63 Penilia avirostris Vol.1, 184-185
Lysmata sp. Vol.2, 151 Periclimenes sp. Vol.2, 148
Petalotrichidae Vol.1, 78
Macrophthalmus sp. Vol.2, 167 Petalotricha Vol.1, 78
Macrosetella Vol.2, 122 Petalotricha ampulla Vol.1, 78-79
Macrosetella gracilis Vol.2, 122-123 Pilidium Larvae Vol.1, 162-163
Metacylididae Vol.1, 64 Pleopis polyphemoides Vol.1, 186-187
Metacylis Vol.1, 68 Pleurobrachia pileus Vol.1, 154-155

K u wait I n s titute f or Scienti f ic R e s earc h 19 5


Pneumodermatidae gen.sp. Vol.1, 176 Temora turbinata Vol.2, 54-55
Podocoryne sp. Vol.1, 138-139 Thalassocaris obscura Vol.2, 145
Pontellidae Vol.2, 58 Thalia sp. Vol.2, 177
Pontella Vol.2, 74 Thenus orientalis Vol.2, 155-157
Pontella danae Vol.2, 74-75 Tintinnidae Vol.1, 96
Pontella investigatoris Vol.2, 76-77 Tintinnidiidae Vol.1, 14
Pontellopsis Vol.2, 78 Tintinnopsis Vol.1, 20
Pontellopsis herdmani Vol.2, 78-79 Tintinnopsis acuminata Vol.1, 20-21
Protorhabdonella Vol.1, 80 Tintinnopsis ampla Vol.1, 22-23
Protorhabdonella curta Vol.1, 80 Tintinnopsis angusta Vol.1, 24
Protorhabdonella simplex Vol.1, 81 Tintinnopsis baltica Vol.1, 25
Pseudevadne tergestina Vol.1, 188-189 Tintinnopsis beroidea Vol.1, 26
Pseudodiaptomidae Vol.2, 48 Tintinnopsis compressa Vol.1, 27
Pseudodiaptomus Vol.2, 48 Tintinnopsis dadayi Vol.1, 28-29
Pseudodiaptomus arabicus Vol.2, 49 Tintinnopsis directa Vol.1, 30-31
Pseudodiaptomus ardjuna Vol.2, 50 Tintinnopsis failakkaensis Vol.1, 32-33
Tintinnopsis gracilis Vol.1, 34-35
Rhabdonellidae Vol.1, 80 Tintinnopsis karajacensis Vol.1, 36
Rhabdonella Vol.1, 82 Tintinnopsis lobiancoi Vol.1, 37
Rhabdonella conica Vol.1, 82-83 Tintinnopsis lohmannii Vol.1, 38
Rhabdonella sp. Vol.1, 85 Tintinnopsis mortensenii Vol.1, 39
Rhabdonella striata Vol.1, 84 Tintinnopsis nana Vol.1, 40
Rhopalophthalmus sp. Vol.2, 126-127 Tintinnopsis orientalis Vol.1, 41
Rotifera Vol.1, 164-165 Tintinnopsis parva Vol.1, 42
Tintinnopsis parvula Vol.1, 43
Sagitta enflata Vol.2, 169 Tintinnopsis radix Vol.1, 44-45
Sagitta neglecta Vol.2, 170 Tintinnopsis rotundata Vol.1, 46
Sagitta pulchra Vol.2, 171 Tintinnopsis sacculus Vol.1, 47
Sagitta regularis Vol.2, 172 Tintinnopsis schotti Vol.1, 48-49
Salpingacantha Vol.1, 117 Tintinnopsis tocantinensis Vol.1, 50-51
Salpingacantha unguiculata Vol.1, 117 Tintinnopsis turbo Vol.1, 52
Salpingella Vol.1, 114 Tintinnopsis undella Vol.1, 53
Salpingella attenuata Vol.1, 114-115 Tintinnopsis urnula Vol.1, 54-55
Salpingella rotundata Vol.1, 116 Tortanidae Vol.2, 90
Sanderia malayensis Vol.1, 150-151 Tortanus Vol.2, 90
Sapphirinidae Vol.2, 106 Tortanus barbatus Vol.2, 90-91
Sapphirina Vol.2, 110 Tortanus forcipatus Vol.2, 92-93
Sapphirina nigromaculata Vol.2, 110-111 Turbellaria Larvae Vol.1, 160-161
Sepiolidae gen.sp. Vol.1, 177
Sergestes sp. Vol.2, 142-143 Undellidae Vol.1, 94
Solenocera crassicornis Vol.2, 132-133 Undella Vol.1, 94
Solenocera hextii Vol.2, 134 Undella dilatata Vol.1, 94-95
Solmundella bitentaculata Vol.1, 142-143 Upogebia sp. Vol.2, 154
Stenopus sp. Vol.2, 144
Subeucalanus Vol.2, 29 Weelia cylindrica Vol.2, 176
Subeucalanus flemingeri Vol.2, 30-31
Subeucalanus subcrassus Vol.2, 32 Xystonellidae Vol.1, 86
Synchaetidae Vol.1, 164-165 Xystonella Vol.1, 87
Xystonella treforti Vol.1, 87
Tadpole Larva Vol.2, 178 Xystonellopsis Vol.1, 88
Temoridae Vol.2, 51 Xystonellopsis gaussi Vol.1, 88
Temora Vol.2, 51
Temora discaudata Vol.2, 52-53 Zoothamnium sp. Vol.1, 120-121

19 6 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f


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